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Volume 24 V, ma as 1970 Number 1
De RMN

JOURNAL,

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

26 March 1970

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

D. F. Harpwick, Editor of the Journal
C. V. Covet, Editor of the News
S. A. Hesse., Manager of the Memoirs

EXECUTIVE COUNCIL

E. B. Forp (Oxford, England ), President

C. L. Remincron (New Haven, Conn. ), President-elect
L. P. Brower (Amherst, Mass.), Ist Vice President

L. M. Martin (Prescott, Ariz.), Vice President

J. W. TiwpvEN (San Jose, Calif.), Vice President

S. S. Nicotay (Virginia Beach, Va.), Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

Members at large (three year term): W. C. McGurrin (Ottawa, Ont.) 1971
J. F. G. Cuarxe (Washington, D.C.) 1970 Y. Nexrurenxo (Kiev, U.S.S.R.) 1971
H. K. Cuencu (Pittsburgh, Penna.) 1970 B. Matuer (Jackson, Miss.) 1972

B. Wricut (Halifax, N.S.) 1970 M. Ocata (Osaka, Japan) 1972

A. E. Brower (Augusta, Me.) 1971 E. C. Wextuinc (Merida, Mexico) 1972

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
and the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

Membership in the Society is open to all persons interested in the study of
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’
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Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests.
In alternate years a list of members of the Society is issued, with addresses and
special interests. There are four numbers in each volume of the Journal, scheduled
for February, May, August and November, and eight numbers of the News each year.

Active members—annual dues $8.00
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Send remittances, payable to The Lepidopterists’ Society, and address changes
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The Lepidopterists’ Society is a non-profit, scientific organization. The office of
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520.
Second class postage paid at Lawrence, Kansas, U.S.A. 66044.

JOURNAL OF

Tue LerrporprTreRIsts’ SOCIETY

Volume 24 1970 Number 1

NEW RECORDS FOR NEW JERSEY WITH NOTES OF OTHER
SCARCE CAPTURES IN 1967

Jos—EPpH MULLER
R.D. #1, Lebanon, New Jersey

Extensive aerial spraying against gypsy moths and mosquitoes, as well
as air pollution may have been instrumental in an apparent sharp decline
of Lepidoptera and most other orders of insects in New Jersey during
1968. Hundreds of blooming milkweed (Asclepias) did not attract a
single butterfly in Cape May. Just as poor was blacklight and bait
collecting in different localities in the State. Many larvae, not accepting
their favorite food, died. This, my poorest collecting season in 25 years,
induced me to write this paper.

In 1967, fourteen species not previously taken by the author in New
Jersey were collected; these included seven species new for the State.
Following is an annotated list of the most interesting captures.

SATYRIDAE

Euptychia mitchellii (French). A series was collected near Newton,
July 13-19. These dates are a few days later than in previous years.
Though similar habitats have been investigated in northern New Jersey,
no other colony has been discovered.

Lethe eurydice eurydice (Johanson). A few years ago F. Rutkowski
discovered this species in a wet meadow near Newton. This season
a series was collected by Rutkowski and the author, July 13-19. In-
dividuals from this population are pale, with eyespots small and the
postmedian band on the underside serrate and irregular. About a mile
from this meadow is a wooded swamp where the darker subspecies, L. e.
appalachia Chermock, flies. This subspecies has darker upper- and
undersides, larger eyespots, and a rounded instead of serrate and irregular
postmedian band on the underside. Each subspecies seems to be limited

2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

to its own slightly different habitat. The latter subspecies has not
previously been reported from the State.
Cercyonis pegala alope, form “maritima” (Edwards). Lakehurst, July 4.

LYCAENIDAE

Euristrymon ontario ontario (Edwards). To my knowledge not more than
six E. ontario had been collected in New Jersey before 1967. After the
discovery of a new area at Lakehurst with dogbane (Apocynum) in
bloom, 17 specimens were caught on July 1 and 4 by several collectors.
Most of the hairstreaks were netted on dogbane, although milkweed
(Asclepias) was plentiful. This area is surrounded by several species
of oak (Quercus), wild cherry (Prunus), and crab apple (Crataegus).
In the hope of obtaining eggs, five females of ontario were kept alive for
several weeks, with dogbane flowers and sugarwater provided for
food. Although the butterflies were confined with oak, wild cherry, and
crab apple, all died without ovipositing.

Satyrium calanus falacer, ab. “heathii” (Fletcher). In the same meadow
where Thymelicus lineola form “pallida” and Euristrymon ontario were
found, one individual of S. c. falacer, ab. “heathii” was collected by F.
Rutkowski on July 1. This specimen shows a bluish border on the
underside of both fore and hindwings. This form has not been reported
from New Jersey previously.

Satyrium liparops strigosus (Harris). Three individuals with orange-
brown patches on the upperside of the forewings were collected at
Lakehurst on July 4 by F. Rutkowski and the author. These were netted
along the railroad tracks where only a few plants of dogbane were
growing. The blossoms attracted mostly S. lL. strigosus, while other
hairstreaks were scarce.

Lycaena phlaeas americana, ab. “fasciata” Strecker. In a large meadow
near Cassville, F. Rutkowski and the author found this species very
numerous. One aberrant individual, with elongated spots, was discovered
among several hundred which were examined.

HESPERIIDAE

Thymelicus lineola, form “pallida” Tutt. Although this introduced species
is widely distributed throughout New Jersey, the pale form “pallida”
was not recorded here before J. B. Ziegler and the author caught eight
specimens at Lakehurst on July 1. Four more specimens were taken at
the same locality on July 4, and more were seen.

VOLUME 24, NUMBER 1 3

NOCTUIDAE

Acronicta rubricoma Guenée. Black light; Lakehurst, June 4.

Acronicta dactylina Grote, melanic form. Black light; Lebanon, June 27.
A new record for the State.

Eurois occulta Linnaeus. Black light; Montague, August 27.
Oncocnemis saundersiana Grote. Black light; Lebanon, October 28.
Agrotis buchholzi Barnes & Benjamin. Black light, Lakehurst, June 4.
Eupsilia morrisoni Grote. Black light; Lebanon, November 18 and 23.

Neperigea costa Barnes & Benjamin. Black light; Montague, July 27.
A new record for the State.

Magusa orbifera, “divaricata” Grote. Black light, Lebanon, August 21.

Amolita roseola Smith. Black light; Montague, July 30. A new record
for the State.

Abrostola urentis Guenée. Black light; Montague, October 10.

Catocala maestosa Hulst. Bait trap; Lebanon, August 28. A new record
for the State.

Zale phaeocapna Franclemont. Black light; Lebanon, April 27. Deter-
mined by genitalic dissection. A new record for the State.

Zale metatoides McDunnough. Black light; Montague, June 10. A new
record for the State.

Gabara pulverosalis Walker. Black light; Lakehurst August 14. A new
record for the State.

Rivula propingalis Guenée. Black light; Lebanon, October 28.
I wish to thank C. F. dos Passos and A. E. Brower for determining some

of the specimens.

A NEW SUBSPECIES OF BREPHIDIUM EXILIS FROM YUCATAN
(LEPIDOPTERA: LYCAENIDAE )

Harry K. CLENCH
Carnegie Museum, Pittsburgh, Pennsylvania 15213

Some years ago Eduardo C. Welling, of Mérida, Yucatan, México,
sent me a few specimens of a Brephidium he had taken on the north
coast of Yucatan. It was obvious, as soon as they had been examined
genitalically, that they represented exilis Boisduval, but they belonged to

4 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Brephidium exilis yucateca Clench. Top left, male holotype, upperside;
top right, same specimen, underside. Bottom left, female paratype, Progreso, Yucatan,
15.x.1958, upperside; bottom right, same specimen, underside.

no known subspecies. Publication was deferred, partly in the hope of
obtaining additional material, and partly because more information was
desirable on the distribution of exilis, particularly along the eastern coast
of México. Both of these desiderata have now been filled.

Brephidium exilis yucateca Clench, new subspecies

This new subspecies differs strikingly from both nominate exilis
Boisduval and subspecies isophthalma Herrich-Schaffer. It is about the
size of B. e. exilis and therefore slightly larger than B. e. isophthalma.
Above as dark as isophthalma and darker than, and without golden sheen
of nominate exilis.

Hindwing often with row of faint pale (ashen) lunules capping subterminal black
spots. Underside of forewing with two sharp and distinct, parallel rows of pale
subterminal bars, heavier than pm series (in both e. exilis and isophthalma these
usually are fainter than pm series), basal area with two sharp and clear, pale post-
basal ring-like marks, one in cell and one in base of Cuz-2A below cell, (nearly
always absent in nominate exilis; in isophthalma sometimes absent and some-
times present, yet faint, the variability depending, at least to some extent, on the
island source). Pm series of forewing below sharper, more distinct, with component
bars slightly thicker than in either of other two subspecies; pm area of forewing
underside without orange, ground color and dark hindwing discal markings chocolate
brown, often grayish, very nearly concolorous with the forewing base (in nominate
exilis flushed with golden orange, contrasting sharply with the gray basal area,
this golden orange extending over even to the discal brown markings on hindwing;
similar in isophthalma, the orange duller and less extensive ).

Holotype male México, Progreso, Yucatan, 15 October 1958 (E. C.
Welling ); ( ¢ genitalia slide no. C-824, CM ). Three paratypes, same data;
seven paratypes, same locality and collector, XII-3-1962; six paratypes,

VOLUME 24, NUMBER 1 5

Chicxulub Puerto [about 2 km E of Progreso], Yucatan, I-8-1967 (E. C.
Welling). All specimens deposited in Camegie Museum, type series
no. 516.

Nominate Brephidium exilis has been found over an extremely large
area, from Nebraska to Oregon, south in the east to Veracruz and in
the west through Baja California, Sonora and Sinaloa. Godman & Salvin
(1887, Biol. Centr.-Amer. Rhop. 2: 109) also record it from two salt
marshes on the Pacific coast of Guatemala. These specimens should be
re-examined to ascertain their subspecific identity.

Large portions of this area are probably inhabited only on a transient
basis, for B. e. exilis is a great wanderer. It is doubtful that it can over-
winter north of central Texas, or above 3-5000 feet elevation southward.
The primary residence areas are in coastal salt flats and salt marshes
in arid or semi-arid parts of the Lower Sonoran, Subtropical and Tropical
life zones. To some extent, particularly from western Texas to California
and probably south into Chihuahua and Coahuila, it may also be a per-
manent resident in interior desert areas.

When Lee D. Miller and I went to eastern Mexico in 1966 one of
our objectives was to learn as much as we could about the distribution
of B. exilis in that region. Between the Brownsville area of southern
Texas, where nominate B. exilis occurs, and Progreso, Yucatan, where
yucateca is found, lie roughly a thousand miles of coast from which we
knew of not a single record of the species. We surveyed coastal areas
in as many places as time and accessibility allowed: in the vicinity of
Tampico, on the Tamaulipas-Veracruz border; the long stretch from
Tecolutla to Cardel; another stretch from near the city of Veracruz south
to beyond Alvarado; and the vicinity of Coatzacoalcos. A few years
previously Gary N. Ross had visited the coast near the Tuxtla Mountains
to search for B. exilis at my request. The results, with one exception,
were everywhere the same, no evilis was found, and no suitable environ-
ment for it. The one exception was a small area 16 miles north of
Cardel where we found a few salt flats, in one of which we collected on
January 19, 1966 and found a single individual of nominate B. exilis.
This area is an enclave of semi-arid conditions (desert scrub and thorn
scrub on the hills back of the flats) and B. exilis is undoubtedly resident
here. This locality is widely disjunct from the next suitable area to the
north (probably somewhere on the coast of Tamaulipas north of
Tampico!). There is also a considerable distance between this colony

1 On our trip we tried to reach this coastal region east of Ciudad Victoria. We got as far
as Soto la Marina but were advised that the road from there to the coast, about 30 miles, was
impassible.

6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

of nominate B. exilis and the nearest known colony of B. e. yucateca,
which is probably a real distributional gap as the intervening area is too
mesic in character.

B. exilis yucateca is most likely restricted to the north coast of Yucatan,
another enclave of arid to semi-arid conditions. The new subspecies
seems to be most closely related to B. e. isophthalma (known from
the Bahamas, Cuba, Hispaniola and Jamaica). In many ways B. e.
isophthalma is intermediate between B. e. exilis and B. e. yucateca.

Godman & Salvin mention some specimens of exilis from Venezuela,
and this record is repeated by Draudt (1921, in Seitz, Grossschmett. Erde
5: 820). There are two specimens in the Carnegie Museum that may
possibly be from Venezuela. They come from the Holland Collection,
and Holland obtained them from Staudinger. They bear no locality data,
but the same style of labelling occurs on some other material of probable
Venezuelan origin. In any event these two specimens seem to represent
a new subspecies, nearer to B. e. yucateca than to either B. e. exilis or
B. e. isophthalma.

AN INTERESTING NEW SPECIES OF THE NEW WORLD
GNORIMOSCHEMINI (GELECHIIDAE) FROM THE
LESSER ANTILLES

Daxror F. PovoLny

Department of Zoology, College of Agriculture, Brno, Czechoslovakia

Several years ago, I had the opportunity of studying interesting ma-
terial of the tribe Gnorimoschemini in the Naturhistorisches Museum in
Vienna. Among their series of this difficult gelechiid group I discovered
a small form which I could not identify. After later study of numerous
American species (Povolny, 1967) and a detailed consideration of this
specimen I decided to describe this moth as a new species belonging
to the genus Keiferia Busck, 1939.

Keiferia rusposoria Povolny, new species
In general appearance this is a small brownish moth, having the fore-
wings dusted with minute darkish scales, forming an indistinct pattern.
Labial palpus not too slender, lacking scales on second segment, its outer surface

markedly grey spotted, inner surface nearly uniform pale cream, third segment pale
with two (one basal and one subterminal) rings of dark grey scales. Frons bright,

VoLUME 24, NUMBER | 7

Fig. 1. Pattern of the forewing of Keiferia rusposoria Povolny.

patagia brownish. Antennal segments brown with distinct dark rings. Forewing
(fig. 1) superficially with ground colour brown; under low magnification individual

scales light to brown, mixed with single scales having dark brown to black tips;
irregular groups of these dark scales forming an indistinct pattern of dark dots
disseminated irregularly over wing; marginal dots around apex more distinct (these
dots characteristic of the tribe). Hindwing medium-grey with fine fringe scales;
costal margin near base with a group of long, brightly coloured bristles, nearly half
as long as costa. Underside of coxae and femora pale cream, nearly uniform, with
only isolated dark scales. Tibiae and tarsi markedly bright and dark spotted to
ringed.

Length of forewing, 3.8 mm.

Male genitalia (fig. 2) characteristic of the genus Keiferia with the heavy thorn-
like process on uncus, which is seldom found elsewhere in members of the
Gnorimoschemini. Other structures of the male genitalia, especially the sacculus
and its processes, also demonstrate the close relationship between the new species
and Keiferia lycopersicella (Walsingham ).

Holotype male: West Indies, Grenada, Balthazar (Windward side),
(H. H. Smith) Walsingham Collection, 1910-427 65173. Deposited in
Naturhistorisches Museum in Vienna.

Keiferia rusposoria may be best distinguished from the two other mem-
bers of the genus, K. lycopersicella and the very similar K. elmorei
(Keifer) by differences in the male genitalia (figs. 2, 3). The thorn-like
process of the uncus is heavier in the new species than in K. lycopersicella,
being only a little shorter than the tips of valvae. In K. rusposoria the
valvae are straight, not S-shaped. The paired processes of the sacculus
are present, but the medial pair is tiny with its branches separated by a
shallow indentation, whereas the other pair of saccular processes is very
long and slender. The saccus is substantially shorter than that of K.
lycopersicella, being comparatively thick with the tip rounded. The
aedeagus is also visibly different from that of K. lycopersicella, being
shorter without the characteristic inflation of the caecum aedeagi, and
moderately curved.

As for the taxonomic position of K. rusposoria, it appears clear that the
species is congeneric with K. lycopersicella but represents a second
distinctive morphotype of the genus. From the polyphagous species K.

lycopersicella, Keifer (1936) distinguished a slightly different species,

8 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 2. Male genitalia of Keiferia rusposoria Povolny, Holotype, Balthazar,
Grenada, West Indies.

Keiferia elmorei. The separation of this sibling from K. lycopersicella
is based on its specialization to indigenous Solanum species in California.

Keiferia rusposoria was collected on Grenada, Lesser Antilles, and thus
might represent a species originating by long-standing geographic isola-
tion. The native distribution of K. lycopersicella, which is referred to as
the “tomato pinworm,” is not known; as the species has been secondarily
introduced as a pest to many parts of Central and South America, and
the West Indies, where it perhaps originally did not occur.

The discovery of K. rusposoria is therefore of high interest, as it is a

VoLUME 24, NUMBER 1 9

Fig. 3. Male genitalia of Keiferia lycopersicella (Walsingham, 1897), Lectotype
(British Museum (Natural History) London) St. Croix, Dan. West Indies.

very distinct species of a genus, which appeared to be monomorphic so
EME

According to Jerry A. Powell, University of California, Berkeley, to
whom I feel greatly obliged for his cooperation in revision of the
manuscript, H. H. Smith collected important material of “microlepidop-
tera” for the British Museum (Natural History) in various parts of the
New World before and after the turn of the century. The windward
side of the small island Grenada in the Lesser Antilles is the east-facing
side.

10 JourNAL OF THE LEPIDOPTERISTS SOCIETY

The specimen might have been sent by Walsingham to Rebel from
London to Vienna, because Rebel was dealing also with the former
collective genus “Lita auct.,” to which this species belonged.

LITERATURE CITED

Kemer, H. H., 1936. California Microlepidoptera. X. Mo. Bull. Calif. Dept.
Agric., 25: 349-359.

Povotny, D., 1967. Genitalia of some Nearctic and Neotropic members of the
tribe Gnorimoschemini (Lep., Gel.). Acta ent. Mus. Nat. Pragae, 37: 51-126.

A NEW PERISAMA (NYMPHALIDAE: CALLICORINI) FROM
BOLIVIA

Joun H. Masters!
Box 7511, St. Paul, Minnesota

During 1962-63, Franz Steinbach, of Cochabamba, Bolivia, consigned
to me 395 specimens of Perisama Doubleday (including Orophila Staud-
inger) collected near Alto Palmar, Chapare Province, Bolivia. The
“Perisama comnena” in this group were particularly interesting because
they apparently included two distinct phenotypes; one characterized by
the normal large red basal patch on the ventral surface of the forewing,
the other without it. This is the type of variation that is normally as-
sociated with geographical subspecies; however, both forms were sympatric
at Alto Palmar. An examination of the male genitalia demonstrated that
the two “forms” were in reality distinct species and, after a canvass of
the literature, I am of the opinion that the second species is heretofore
unnamed.

Perisama rusea Masters, new species

Mae: Upperside of both wings deep-black ground; iridescent green rays extending
outwardly from base of forewing, along base of discal cell and vein 2V, until they
almost meet a diagonal iridescent green band, 2 to 3 mm wide, which crosses the
wing from costal margin, at end of cell to inner margin at tornus; one (occasionally
two) iridescent green spot(s) in subapical area, near costal margin. Hindwing
with an iridescent green band, 2mm wide, extending through limbal area parallel
to outer margin.

Underside of forewing with dull black ground color except for a golden apex;

base of wing with a small golden patch, confined primarily to cell and not over 8 mm
long, having a tint of red at its outer margin; from this golden patch, a blue-green
streak extends along costal margin of cell to a white triangular patch at end of cell

which in turn precedes three small blue spots, in cells M2, M3 and Cui, which form

| Research Associate, Section of Insects and Spiders, Carnegie Museum, Pittsburgh.

VOLUME 24, NUMBER | Wt

y \)
Ea Sy

Cc

Figure 1. Male genitalia of Perisama rusea Masters: (A) left lateral view of
genital capsule with inside surfaces shaded; aedeagus (B) removed and_ the
gonostatumen (C) detached and rotated 90° to view ventrally. Male genitalia of
Perisama comnena Hewitson: (D) left lateral view of uncus and tegumen, (E)
aedeagus and (F) ventral view of gonostatumen.

a diagonal line towards tornus. Underside of hindwing a uniform golden base color;
two wavy black lines extending from costal margin, where they are at least 2 mm
apart, to vannal veins where they stop short of anal margin (the eminens pattern,
Schwanwitsch, 1930) with no trace of vestigial eyespots between them.

Male genitalia (figue 1A-C) very distinct from allied species; aedeagus relatively
short (approximately 2 mm long) and not over 75% as long as gonostatumen
(Dillon, 1948) which is distinctly heart shaped anteriorally.

FEMALE: Similar to male but larger and markings for most part, bolder but with
duller colors. The eminens lines, on hindwing underside, meet on costal margin
and between them five golden-brown eyespots, in cells Ms, Me, discal, Cu: and Cup,
faintly visible.

LENGTH OF FoREWwING: Male holotype, 21 mm; female allotype, 23 mm.

Holotype male and allotype female: Bolivia, Alto Palmar, Chapare
Province, District of Cochabamba, 1,100 meters elevation, March 1963.
Types deposited in Carnegie Museum, Pittsburgh.

Paratypes: 22 males from the same locality, various dates, 1962 and
1963. Single paratypes are being placed in the American Museum of
Natural History, New York; The U.S. National Museum, Washington;
The Museum of Comparative Zoology, Harvard, Cambridge, Massachu-

12 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figure 2. Perisama rusea new species, paratype male, Alto Palmar, Chapare,
Bolivia, March 1963: (G) upperside and (H) underside. Perisama comnena
Hewitson, Alto Palmar, Chapare, Bolivia, April 1963: (J) upperside and (K) under-
side.

setts; and the Reading Public Museum, Reading, Pennsylvania. The
remainder are being retained in the author's collection.

Perisama rusea (figure 2G-H) is immediately distinguishable from
Perisama comnena Hewitson (figure 2J-K) by the lack of the large red
basal spot on the ventral surface of the forewings; from Perisama xanthica
Hewitson by the presence of the green limbal band on the dorsal surface
of the hindwing and by a much smaller golden patch at the base of the
ventral surface of the forewing; and from Perisama humboldtii Guer.
by the lack of punctiform eyespots on the ventral surface of the hindwing
and a large red basal spot on the ventral surface of the forewing. All four
of these species are sympatric at Alto Palmar, Bolivia and all have distinc-
tive male genitalia. Examination of several series of P. comnena from Peru
did not reveal any P. rusea among them.

Examination of the male genitalia in Perisama and related genera
(Callicorini) reveals that the uncus, tegumen, saccus, aedeagus and

VOLUME 24, NUMBER 1 eS

gonostatumen all have diagnostic characters at the species level. It is
suprising that previous use has not been made of them. In his revision
of the Callicorini (as Catagrammini), Dillon (1948) characterized the
male genitalia in his definition of the tribe, but did not use them to
diagnose species. The gonostatumen is an interesting structure that is
found only in the Callicorini, including Callithea Feisthamel. It was
named by Dillon (1948) who believed it to be a modification of the
eighth sternite and to function as a prop for the usual genital organs.

LITERATURE CITED

Ditton, L. S., 1948. The tribe Catagrammini (Lepidoptera: Nymphalidae).
Part I. The genus Catagramma and allies. Reading Public Museum Scientific
Publ., No. 8: 1-113.

ScuwanwitscuH, B. N., 1930. Studies upon the wing-pattern of Catagramma and
related genera of South American nymphalid butterflies. Trans. Zool. Soc.
London, 21: 105-284.

MULTIPLE CAPTURE OF CARIA INO MELICERTA
(RIODINIDAE) AT LIGHT

-LeE D. MILLER
The Allyn Foundation, 222 West Adams Bldg., Chicago, Illinois

A rather extensive bibliography has accumulated concerning butter-
flies that have been attracted to light. Most recent records have been of
captures at ultraviolet light, such as reported by Throne (1961) and
Welling (1963). Most of these records involve single specimens, but
there have been a few instances of multiple captures of butterflies
(Donahue, 1962; Phillips, 1962; Welling, 1963; and Hessel, 1965), for
example. Some of the collections have been at ultraviolet light, some at
fluorescent and a few at incandescent light.

There seem to be two explanations for the attraction of butterflies to
light: (1) the butterflies were at rest, were disturbed and these flew
to the light source, and (2) the butterflies are crepuscular species and
were attracted to the light as are many crepuscular Sphingidae and
other moths. Most of the records fall into the first category, and they
almost always are based on one or two specimens; the majority of the
captures I have made at light are in this class, both in the United States
and in the Neotropics. Several of the multiple captures, especially those
of Opsiphanes (Welling, 1963) and Melanitis (Donahue, 1962), are of

14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

crepuscular species, and the captures of Hypaurotis reported by Hessel
(1965) involve a species that is definitely not heliophilic during midday
hours and continues activity into evening (e.g., Chambers, 1963).

During July and August, 1966, I taught a travelling field biology course,
and the students and I often trapped moths at ultraviolet light as part of
the activities without taking any butterflies. The evening of August 2nd
we set up the light at Bentsen-Rio Grande State Park, Hidalgo County,
Texas, on a hot (ca. 85° F.), muggy night, and collecting was excellent
for moths and other insects. The effectiveness of the ultraviolet lamp
was attested to by the fact that very few insects were attracted to the
Coleman lantern we used as a camp light about a hundred feet from the
moth sheet. I was surprised, therefore, to see a specimen of Caria ino
melicerta Schaus, a riodinid, flop down on the table next to the lantern
and begin walking slowly over the surface of the tabletop gently opening
and half closing its wings until I bottled it. A short time later one of the
students took a second specimen at the Coleman lantern, and still later
I captured two more specimens that were behaving in the same manner
as the first. Another of the students also collected a specimen at an
incandescent light in one of the park bath houses. All five melicerta
were taken between 9:00 and 11:00 P.M., C.D.T. A check of the insects
collected at the ultraviolet light revealed no butterflies, even though
hundreds of moths had been taken.

I collected only a single female of C. i. melicerta in two days of col-
lecting in the same area in which the five specimens were collected at
light. This specimen was netted in deep woods resting under a leaf,
and it looked as though this species was crepuscular, or at least helio-
phobic. Two hours’ collecting at Bentsen park in September, 1967, yielded
about fifty specimens at flowers in the brightest sunlight, so C. i.
melicerta is not consistently crepuscular or heliophobic.

These records of this butterfly, I believe, are the first to demonstrate
any butterfly making a “choice” between visible and ultraviolet light,
and it is intriguing that these insects selected the visible light source.
[ have been unable to obtain spectroscopic analyses of the light from a
Coleman lantern, but I suspect that the ultraviolet component is minor,
as is that of an incandescent light. This demonstrates a phenomenom
that is well known, but seldom stated: some insects that will come to
visible light are not attracted to ultraviolet light. The advent of ultra-
violet light as a collecting aid has greatly increased the numbers of both
individuals and species that may be taken during a night’s collecting, but
many species of Lepidoptera cannot be obtained by this method.

VOLUME 24, NUMBER 1 15

LITERATURE CITED

CHAMBERS, D. S., 1963. Evening mating in Hypaurotis crysalus (Lycaenidae).
J. Lepid. Soc., 16: 200 (“1962”).

DonauuE, J. P., 1962. Observations and records of butterflies attracted to light
in India. J. Lepid. Soc., 16: 131-135.

HeEssEx, J. H., 1965. Multiple capture of Hypaurotis crysalus at light. J. Lepid.
Soc., 19: 55-56.

Puitures, L. S., 1962. Nymphalis j-album captured at fluorescent light in Chicago.
ieleeprdeisoc:, 5: LO]:

TuroneE, A. L., 1961. Lycaenopsis pseudargiolus in light trap. J. Lepid. Soc., 14:
DAD:

Wexunc, E. C., 1963. Rhopalocera attracted by ultraviolet light in Central
America. J. Lepid. Soc., 17: 37-38.

BIONOMIC NOTES ON HAETERINI AND BIINI IN
VENEZUELA (SATYRIDAE)

Joun H. Masters?
P.O. Box 7511, St. Paul, Minnesota

HAETERINI

Four genera, Pierella Westwood, Haetera Fabricius, Dulcedo d’Almeida
and Cithearias Hubner (= Callitaera Butler), comprise the Haeterini,
the most primitive tribe of Satyridae. All of the Haeterini have distinc-
tive wing shapes with short, narrow forewings and seemingly enlarged
hindwings. Haetera, Cithearias and Dulcedo are the only clear-winged
Satyridae (see Figure 1, Cithearias andromeda Fabricius). All of the
known species are Neotropical and are found on the forest floors of dense
tropical and subtropical forests.

Virtually nothing has been published concerning the habitat and habits
of these butterflies. I have found only brief notes by Weymer (in: Seitz,
1909) and Brown (1942). Both mention that these satyrids inhabit
forested areas, fly near the ground and are difficult to see or follow in
flight. Weymer mentions that members of the genus Cithearias ( given as
Callitaera) preter the early morning hours for flight.

After having the opportunity to observe several of these species in
Venezuela (1965 and 1966 in Bolivar and 1968 in Barinas), I can add
somewhat to these statements. Determinations of my specimens were
made by Michael Clifton of the British Museum (Natural History) who
is currently preparing a taxonomic revision of the tribe.

The species that I observed seemed to be strictly limited to denser

1 Research Associate, Section of Insects and Spiders, Camegie Museum, Pittsburgh.

16 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figure 1. Cithearias andromeda Fabricius. 88 kilometers south of El Dorado,
Bolivar, Venezuela; 20 Feb. 1966 (J.H.M.) Natural Size.

selva (low and tropical forests) where they flew close to the forest floor
and nearly always lit on the ground. They were distinctly matinal and
crepuscular in habit, flying during both early morning and evening hours.
However, Pierella species, and to a lesser extent the others, would fly
throughout the day during inclement weather. All of the species were
attracted to fruit, but to catch them at it was difficult. Normally fruit
bait is placed in the center of a small area from which all leaves and
underbrush have been cleared and then those species that are attracted,
including Taygetis, Euptychia, Caligo, Morpho, etc., are easily netted.
Haetera and Cithearias species are attracted to fruit but will not alight
in a cleared patch of ground but will alight around its edge instead
where they can be perfectly camouflaged. Of course, if the bait is placed
on uncleared ground, they blend into the background so well that they
are completely inconspicuous while visiting it. The use of bait traps
would seem to be the answer (for a description of bait traps and their
use see Rydon, 1964), but the effectiveness of most traps depends upon
butterflies to fly upward when disturbed. While most butterflies do fly
upward when disturbed, the Haeterini fly horizontally and close to the
ground and thus cannot be taken in any ordinary sort of trap. The best
method of collecting them is to spread fruit on the ground to attract
them into the area, and to then walk through the tract at dusk flushing
them and carefully stalking them.

Haetera piera piera ( Linnaeus )

This species was encountered in heavy selva in both Bolivar and
Barinas. H. piera, with transparent wings just shaded with yellow,
appears ghost-like in flight, just a flicker of yellow moving across the

VOLUME 24, NUMBER 1 IZ

leaves. They fly slowly and alight often, but it is difficult for the eye to
follow them in flight and, unless you actually see them alight, they are
virtually impossible to detect on the ground. They were usually en-
countered as solitary individuals, but the presence of fruit (natural or
bait) on the forest floor would concentrate them into a given area. In
1966, a single male of piera was taken at heliotrope (Masters, 1968), a
bait normally employed to attract Ithomiidae. This might be another
source of evidence of the close phylogenetic relationship of these primi-
tive Satyridae to the Ithomiidae, a relationship pointed out by Fox
(1956). However rather than being attracted to the heliotrope the satyr
was probably attracted to traces of fruit on it as heliotrope and fruit
baits had been spread along the trail at the same time.

Cithearias andromeda Fabricius

Cithearias andromeda (=esmeralda Dbl.) was found in the same
habitat in Bolivar as H. piera but was much scarcer and less apt to ven-
ture into an open area. C. andromeda (figure 1) also has transparent
wings, but shaded with purple instead of yellow, a combination that
makes it even more difficult to observe and follow in the selva. Its flight
is also slow, and apparently its primary defence against predation is the
inconspicuous character of nearly colorless, transparent wings.

Pierella astyoche Erichson

This species was encountered in heavy selva in Bolivar but was much
more prone to come out along trails or to enter small clearings, thus
being much more in evidence and more frequently encountered than H.
piera or C. andromeda. Pierella astyoche is a quite active flier and males
are fond of flying quickly up and down a forest trail close to the ground.
Each male seemed to have its own section of path to patrol and this
activity suggests a form of reconnaissance for females. Interest in patrol-
ling decreased when fruit was spread on the trail and astyoche was less
shy in coming to bait than the clear-winged species. The best method of
capturing them, however, was to note the path of a patrolling male and
to then wait along it to intercept the butterfly in flight. Peak flight for
P. astyoche was from late afternoon until dusk, but a few individuals
seemed to fly throughout the day. On cloudy days they would fly in
numbers all day long, and, along with Taygetis species, would be virtually
the only butterflies encountered on the forest floor.

Pierella hyalinus hyalinus Gmelin

Pierella hyalinus is a large attractive species with deep blue on the

18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

hindwings. P. hyalinus is sympatric with astyoche in Bolivar and has
similar habits, but the two could be readily distinguished on the wing
because of the large size and blue of the rarer hyalinus.

Bint

A single unique species, Bia actorion (L) (= actoriaena Hubner),
comprises the Biini.

Bia actorion actorion ( Linnaeus )

The geographical range of Bia actorion is limited to the lowland rain
forests of the Amazon and Orinoco basins. I found the specific habitat
preference of B. actorion to be those areas where enough sun penetrates
to the forest floor to provide a thick and heavy undergrowth rather than
those areas where the canopy is complete and prevents enough sunlight
to reach the ground to allow for much undergrowth. Solitary individuals
of B. actorion were encountered along trails or in clearings where they
would alight about three feet above the ground among underbrush.
They quickly fly to the base of a bush when alarmed where they are
either perfectly concealed or impossible to get at with a net. Bia actorion
is very striking during flight because of the nearly iridescent upper
wings that flash in the sunlight, giving the appearance of a miniature
Morpho. When the insect alights, the flashing blue suddenly disappears,
which undoubtedly adds to the deception as the cryptic undersides blend
into the background.

LITERATURE CITED

Brown, F. M., 1942. Notes on Ecuadorian Butterflies, V. J. New York Ent. Soc.,
50(4 2 809=380.

Fox, R. M., 1956. A monograph of the Ithomiidae (Lepidoptera). Part 1. Bull.
American Mus. Nat. Hist., 111: 1-76.

Masters, J. H., 1968. Collecting Ithomiidae with heliotrope. J. Lepid. Soc.,
222) OSGeo}

Rypon, A., 1964. Notes on the use of butterfly traps in East Africa. J. Lepid.
Soc., 17(4): 51-58 (“1963”).

Seitz, A., 1909. Gross-schmetterlinge der Erde, 5. Kernen, Stuttgart.

VOLUME 24, NUMBER | 19

STUDIES ON THE BIOLOGY AND SEASONAL HISTORY OF
POLYDORUS ARISTOLOCHIAE (PAPILIONIDAE )

G. H. Munsutr ann S. A. Moiz
Agric. College & Agric. Research Institute, Tandojam, West Pakistan

Polydorus aristolochiae (Fabricius) is the commonest large tailed,
black butterfly of Indo-Pakistan Subcontinent. At Tandojam, West Paki-
stan the larvae have been found in large numbers on Aritolochia bracteata.
The biology of this insect has been studied only by Ghosh (1914). The
observations recorded by us concerning this species are given in the
present paper. The work was under taken during 1962-63 at Tandojam.

Ecc
Description

The eggs of Polydorus aristolochiae are laid on the underside, very
rarely on the upper surface, of host leaves. A high oviposition rate was ob-
served in the months of August and September and again during March
and April. Up to five eggs were collected from each leaf.

The chorion is covered with a reddish brown, soft gum like substance,
in minute granular masses. The latter are arranged somewhat in undulat-
ing ridges meeting at a point. These give the egg the appearance of a
citron with the long axis slightly shorter and a little inclined towards one
side and never perpendicular to the leaf surface.

Incubation Period

Since the butterflies did not lay eggs in captivity, the incubation period
was determined by recording the maximum period required for hatching
of eggs collected in the field. The time taken for hatching of the eggs is
given in Table 1.

Ghosh (1914) recorded that the incubation period is four to five days
from June to September. The results obtained here are in general agree-
ment with his observations. Between the months of October and March
the incubation period is longer due to lower temperatures.

LARVA

The larva is dark brown, and passes through four instars. The duration
of larval stages was studied in the laboratory. The results are presented
in Table 2.

It can be seen from table 2 that the duration of larval life varies from
14 to 62 days. The minimum duration was for larvae which hatched in
September, 1962. Those hatching in the early part of January, 1963,

20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. INCUBATION PERIOD OF EGGS OF Polydorus aristolochiae
COLLECTED IN THE FIELD

Days re- Estimated Average
quired for incubation monthly
hatching period Temperature
Month of eggs in days (Fahrenheit )
September 2-5 5 Days 75.91
October 3-9 9 Days 69.64
November 2-9 9 Days 73.78
December 2-9 13 Days 60.88
January 8-13 13 Days 60.92
February 4-10 10 Days 69.92
March 3-10 10 Days Tass
April 2-6 6 Days 83.75

passed their development during the two coldest months and, hence,
their development required 62 days.

Ghosh (1914) has described the prolongation of larval life as hiberna-
tion. During our studies at Tandojam, however, larvae continued to
feed during this period.

Pura

The length of the pupal period varied from 10 to 230 days as is shown
in Table 3.

The duration of the pupal stage is not correlated with atmospheric
temperature (Table 1). Ghosh (1914) observed the pupal period to last
from 10 to 303 days, and the present observations are in general agree-
ment. However, Ghosh mentions that the insect undergoes a hibernation
followed by an aestivation. This deduction does not seem to be correct
since adult emergence took place in the coldest period as well as in hot
months. Further, pupae from the same batch required widely varying

TABLE 2. LENGTH OF LARVAL LIFE OF P. aristolochiae

Month in which Larval life
egg hatched (in days )
September 14-15
October YS
November 24-31
December 31-43
January 50-62
February 26-35
March 26=85

April 28-34

VOLUME 24, NUMBER | Zak

TABLE 3. PUPAL PERIOD OF P. aristolochiae

Month in which Pupal period

larva pupated (in days )

August 10
September 13-230
October 13-228
November 19-150
December 36-141
January 44-143
February 14-179
March 13-172
April 12-38

periods before emergence of the adult, and it emergence by later pupating
individuals sometimes occurred while from pupae of an earlier batch
emergence of butterflies had not taken place. Thus, there is great range
of individual variation in the pupal period, without apparent effect of
temperature.

The above conclusion is corroborated by observations on pupae col-
lected in the field. One hundred forty pupae were collected on November
5 and 25, 1962, from a small area at Tandojam which was under constant
observation. Pupation had taken place on 5th and 25th November
respectively.

LIFE CYCLE

According to the observations made on the length of incubation,
larval and pupal periods (Tables 1-3), it is seen that the butterfly

TABLE 4. EMERGENCE OF ADULTS FROM FIELD COLLECTED PUPAE

Date of Month of adult Pupal period
collection emergence (in days )
5 Nov. 1962 February 88-115
March 127-146
April 153-176
May 184-207
June 215-224
July 246-258
August 275
25 Nov. 1962 December 23
January ol
February 12
March 120
April 146

May 166-176

22, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

requires from one to about 9 months to complete its life cycle (exclusive
of adult).

It may, therefore be concluded that the insect may have up to about
7 overlapping generations in a year. Normally, in the field it probably
has fewer.

Field surveys revealed the presence of all the stages on the host plants
throughout the year.

NATURAL ENEMIES

Thompson (1946) reported Apanteles aristolochiae Wlkn. as a parasite
of this butterfly. No parasites of eggs or larvae were found during this
study.

ACKNOWLEDGMENT

To Jerry A. Powell, University of California, Berkeley, we wish to
express our sincere appreciation for correcting and improving the
manuscript.

LITERATURE CITED

Guosu, C. C., 1914. Life histories of Indian Insects. V. Lepidoptera. Mem.
Dept. Agric. India, Ent. Ser. 5, 1: 53-58. Agric. Res. Inst., Delhi, Pusa.
THompson, W. R., 1946. Parasites of Lepidoptera (N-P). A catalogue of parasites

and predators, Sec. 1. Pt. 3. Imperial parasite service. Belleville, Ontario,
Canada.

A NEW CALLITHOMIA (ITHOMIIDAE) FROM BOLIVIA

Joun H. Masters!
P.O. Box 7511, Saint Paul, Minnesota

During 1962-1965 I received several large consignments of Bolivian
[thomiidae from Franz Steinbach of Cochabamba, Bolivia. Perhaps the
most interesting specimen among these is an unique and previously un-
described Callithomia. The late Dr. Richard M. Fox, acknowledged
expert in the Ithomiidae, examined this specimen, recognized it as a new
species and intended eventually to describe it himself. With the death
of Dr. Fox on April 28th, 1968, I have assumed the task of describing
the insect and take pleasure in naming it after him.

Research Associate, Carnegie Museum, Pittsburgh, Pennsylvania.

VoLUME 24, NUMBER 1 23

Figure 1. Callithomia foxi Masters. Holotype male: A, upperside; B, underside.

Callithomia foxi Masters, new species

MALE: Upperside of both wings (figure 1A) transparent with black markings
outlining veins and wing margins. Black markings widening on both wings at costal
and inner margins, wing apices, at end of discal cells and along vein Cu:. Cu on
hindwing and basal parts of veins Cuz and 2V yellow. Inner margin of forewing
sharply concave. Costal margin of hindwing sharply humped at raised oval “Ithomia
patch” from there concave to outer angle.

Underside of wings (figure 1B) a mirror image of upperside except for eight
small white spots in black wing margins—two at apex of forewing (also a dimunitive
in Mz), four at anal angle and two at apex of hindwing. Transparent areas with a
yellowish cast, viewed from below.

Male genitalia figured (figure 2).

LENGTH OF FOREWING: base to tip, 29 mm.

FEMALE: Unknown.

Holotype male: Bolivia, Todos Santos, Prov. Chapare, Dpto. Cocha-
bamba (300 meters elevation), July 1964. Deposited in Carnegie
Museum, Pittsburgh, Pennsylvania.

Dr. Fox (in litt.) believed that this specimen was the only example
of the species in American collections, but noted that there were a couple
of examples resting unnamed in European collections. The subterminal
spine on the aedeagus places Callithomia foxi in that section of the genus

24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figure 2. Callithomia foxi Masters male genital structure. Aedeagus removed
and shown below. Left valva removed and not shown.

that includes xantho (Felder), methonella (Weymer), inturna (Fox),
epidero (Bates), lenea (Cramer), drogheda (Weeks) and zingiber Fox.
Of these seven, C. epidero, methonella, xantho and interna have a strong
resemblance to C. foxi. Callithomia xantho and interna, and to a lesser
extent methonella, are separable by the lack of heavy black scaling along
vein Cu, and the end of the cell, this is especially noticeable on the
hindwings. Callithomia epidero and methonella have red to brown
patches at the anal angle of the hindwing, that are lacking in C. foxi.
None of the species in the other section of the genus have a similar
appearance.

VOLUME 24, NUMBER 1 DAD)

AN OBSERVATION ON THE USE OF COLOR FOR SPECIES-
RECOGNITION IN HELICONIUS BESCKEI (NYMPHALIDAE )

MicHAEL G. EMSLEY
George Mason College, U. of Virginia, Fairfax, Va.

Crane (1955) has shown red to be an important courtship releaser in
Heliconius erato hydara Hewitson, and Swihart (1963, 1964) has con-
firmed the significance of red in that species by neurophysiological
techniques.

During the course of genetical studies on Heliconius, living specimens
of the mimetic pair, Heliconius erato phyllis (Fabricius) and H. besckei
Ménétriés, from Rio de Janeiro, Brazil, were caged with specimens of H.
melpomene melpomene (Linnaeus) from Trinidad. Both H. e. phyllis
and H. besckei have red forewing bands and a yellow hindwing bar,
but H. m. melpomene has only red forewing bands. All colors are on a
black background.

It was observed that male erato phyllis and male melpomene would
each fly towards members of either of the other two species in the cage,
but presumably because secondary courtship releasers were lacking they
would then disengage and fly off (all the females had already been
mated). H. besckei, however, though appearing in good health, would
only pursue the phyllis, not the melpomene. The color difference between
these two is the lack of yellow in melpomene. The courtship releaser is
known to be red in erato and is likely also to be red in melpomene.
Possibly it is yellow in besckei.

Whether the courtship releaser in besckei is red or yellow is an impor-
tant behavioral character which should be considered in deducing the
course of the evolution of mimicry in Heliconius.

LITERATURE CITED

CRANE, J., 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara
Hewitson, with special reference to the social use of color. Zoologica, N. Y., 40:
167-196.

SwiHART, S. L., 1963. The electroretinogram of Heliconius erato (Lepidoptera)
and its possible relation to established behavior patterns. Zoologica, N. Y., 48:
155-165.

1964. The nature of the electrorectinogram of a tropical butterfly. J. Insect
Physiol., 10: 547-562.

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

A REVISED SYNONYMIC CATALOGUE WITH TAXONOMIC
NOTES ON SOME NEARCTIC LYCAENIDAE

Cyrit F. pos Passos
Washington Corners, Mendham, New Jersey

INTRODUCTION

The publication by Harry K. Clench ([{1961]: 177-220) and John
C. Downey ([1961]: 230-242) in Ehrlich & Ehrlich’s How to Know the
Butterflies of revisions of the Theclini by the former and the Plebejini
by the latter rendered another part of the present author's synonymic list
(1964), at least obsolete insofar as the Theclini is concerned. This
situation is similar to the one explained in a previous paper on the
Nearctic Melitaeinae (1969, J. Lepid. Soc., 23: 115-125).

Clench revised the Theclini and in the course of his work proposed
eight new generic or subgeneric names, besides introducing into our
fauna some names that had heretofore been considered Neotropical and
eliminating some that are no longer considered Nearctic. Downey, on
the other hand, merely rearranged the order in which the genera and
some of the species appear in the synonymic list so that it is not con-
sidered necessary to repeat his work, no reasons having been given for
such action. Clench also made no vital changes in this author's
Lycaeninae which he gives tribal rank (Lycaenini) except that he
combines Lycaena cupreus with L. snowi (1961: 222) as one species al-
though recognizing apparently both as good subspecies. Also he does not
clearly recognize the subgenus Tharsalea Scudder, 1876, although he may
recognize it subgenerically in another faunal area, and arranges the
species in Lycaena in somewhat different order. These are subjective
matters with which one cannot quarrel, but in neither case is it deemed
necessary to rewrite those parts of the synonymic list.

Neither Clench’s nor Downey’s works give any synonymies and do not
list categories lower than species. This was done in accordance with the
wish of the authors of How to Know the Butterflies, but is unfortunate
for the student because he cannot tell what has become of subspecies,
lower categories, and synonyms.

The present paper follows the general style and form of the synonymic
list, followed by notes giving explanations for the changes made. An
effort has been made to fit all names into the list by giving numbers to
them starting with 349 where the Lycaenidae start. A similar arrange-
ment of the genera was impossible because of the many changes made
by Clench.

bo
~l

VOLUME 24, NUMBER |

The thorough and badly needed revision of the Theclini by my
colleague Harry K. Clench, Curator of Lepidoptera at the Carnegie
Museum, is of such value and importance that it should be made available
in catalogue form to purchasers of the synonymic list. My thanks are
extended to Mr. Clench for clearing up some points that were uncertain
in his work or not easily understood. He has been most liberal in his
assistance.

I am indebted also to my colleague, F. Martin Brown, for placing at my
disposal a ms on the types of the Lycaenid butterflies described by
William Henry Edwards of which he is the senior author with Paul A.
Opler. Brown generously gave permission to use any of his ideas.

Family LYCAENIDAE
Subfamily THECLINAE

HABRODAIS Scudder, 1876
Type: Thecla grunus Boisduval, 1852
Habrodias McDunnough, 1914 (lapsus calami )

349 grunus ( Boisduval), 1852

a g. grunus (Boisduval), 1852

bg. lorquini Field, 1938

form chloris Field, 1938
c g. herri Field, 1938

HYPAUROTIS Scudder, 1876
Type: Thecla chrysalus Edwards, “1872-3” (1873) (= Thecla crysalus
Edwards, “1872-3” [1873] )
350 erysalus (Edwards ), “1872-3” (1873)
chrysalus Auctorum
chryaslus (Edwards), 1884 (lapsus calami)
form citima (H. Edwards ), 1881

CHLOROSTRYMON Clench, [1961]
Type: Thecla telea Hewitson, 1868
*351 simaethis ( Drury ), “1770” [1773]
a s. sarita (Skinner), 1895
302 telea (Hewitson ), 1868
303 maesites ( Herrich-Schaffer ), 1864

PHAEOSTRYMON Clench, [1961]
Type: Thecla alcestis Edwards, “1870-1” (1871)
304 alcestis (Edwards), “1870-1” (1871)
a a. alcestis (Edwards), “1870-1” (1871)
b_ a. oslari (Dyar), 1904

28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

HARKENCLENUS nom. nov.
pro Chrysophanus Hubner, 1818 (opinion 541, name 1235)
Type: Chrysophanus mopsus Hubner, 1818 (opinion 541, name 1235)
(= Papilio titus Fabricius, 1793) (opinion 541, name 1605)
3559 titus ( Fabricius ), 1793
a t. titus (Fabricius ), 1793
b t. mopsus (Hubner), “1818” [1809-13]
c t. watsoni (Barnes & Benjamin), 1926
d t. immaculosus (Comstock), 1913

SATYRIUM Scudder, 1876
Type: Lycaena fuliginosa Edwards, 1861
Callipsyche Scudder, 1876
Strymon Auctorum (partim )
Thecla Auctorum (partim )
306 fuliginosum (Edwards ), 1861
a f. fuliginosum (Edwards), 1861
suasa ( Boisduval ), 1869
ab. immaculata Gunder, 1927
b_ f. semiluna Klots, 1930
357 ~behrii (Edwards ), “1870-1” (1870)
a hb. behrii (Edwards ), “1870-1” (1870)
kali (Strecker ), “1872” [1878]
ab. nigroinita (Gunder ), 1924
bb. erossi (Field), 1938
c b. columbia (McDunnough), 1944
358 auretorum (Boisduval), 1852
a a. auretorum (Boisduval ), 1852
tacita (H. Edwards ), 1881
b a. spadix (H. Edwards), 1881
359 tetra (Edwards ), “1870-1” (1870)
adenostomatis (H. Edwards ), 1877
360 saepium ( Boisduval), 1852
chalcis Edwards, “1868-9” (1869 )
soepium ( Boisduval ), 1852 (lapsus calami)
a 8. saepium (Boisduval), 1852
form fulvescens (H. Edwards ), 1877
form chlorophora (Watson & Comstock ), 1920
form provo (Watson & Comstock ), 1920
b s. okanagana (McDunnough), 1944

VoLUME 24, NUMBER |

361

362
363

364
365

366

367

368

369

liparops (Le Conte), [27 July 1833]
a I. liparops (Le Conte), [27 July 1833]
b_ I. strigosa (Harris), 1862

ab. pruina (Scudder ), 1889
c |. fletcheri (Michener & dos Passos ), 1942

liparops (Fletcher nec Le Conte [27 July 1833] )

“1903” [1904]
d 1. aliparops (Michener & dos Passos ), 1942
kingi (Klots & Clench), 1952
calanus ( Hubner ), “1806” [1809]

wittfeldii (Edwards ), 1883
a e. calanus ( Hiibner ), “1806” [1809]
b_ e. falacer (Godart), “1819” [1824]

lorata (Grote & Robinson ), 1867

inorata (Grote & Robinson ), 1867

ab. heathii ( Fletcher ), “1903” [1904]
ce e. godarti (Field), 1938
caryaevorus (McDunnough), 1942
edwardsii (Saunders ), 1869

fabricii ( Kirby ), 1871
sylvinus ( Boisduval), 1852
s. sylvinus ( Boisduval), 1852
s. desertorum (Grinnell), 1917
s. itys (Edwards ), 1882
s. putnami (H. Edwards), 1877

putmani Brown, Eff, & Rotger, 1955 (lapsus calami)
californica (Edwards ), 1862

borus ( Boisduval ), 1869

cygnus (Edwards ), “1870-1” (1871)
acadica (Edwards ), 1862

acadia Brown, Eff, & Rotger, 1955 (lapsus calami)

a a. acadica (Edwards), 1862

souhegan (Whitney ), 1868

souhegon (McDunnough), 1938 (lapsus calami)

ab. muskoka (Watson & Comstock ), 1920

ab. swetti (Watson & Comstock ), 1920
b_ a. coolinensis (Watson & Comstock), 1920
c a. montanensis (Watson & Comstock), 1920
d a. watrini (Dufrane), 1939
dryope (Edwards ), “1870-1” (1871)

Oo @ @ &

29

30 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

EUMAEUS Hiibner, “1816” [1819]
Type: Eumaeus minyas Hubner, “1816” [1819] (= Rusticus adolescens
minijas Hubner, “1806” [1809] )
Eumenia Godart, “1819” [1824]
*370 atala (Poey), 1832
a a. florida Rober, 1926
grayi Comstock & Huntington, 1943
371 minyas (Hiibner ), “1816” [1819] (emendatio )
minijas (Hubner ), “1806” [1809]
toxea (Godart ), “1819” [1824]

OENOMAUS Hiibner, “1816” [1819]
Type: Papilio ortygnus (Cramer ), “1782” [1779]
372 ortygnus (Cramer), “1782” [1779]

MINISTRYMON Clench, [1961]
Type: Thecla leda Edwards, 1882
373 ines (Edwards ), 1882
374 leda (Edwards), 1882
375 elytie (Edwards ), 1877
gen. hiem. maevia (Godman & Salvin ), (1887 )

HETEROSMAITIA Clench, 1964
Type: Thecla bourkei Kaye, 1924
376 spurina Hewitson, “1862” [1867]
377 zebina (H. A. Freeman), 1950
CALYCOPIS Scudder, 1876
Type: Rusticus armatus poeas Hubner, “1806” [1811] (= Hesperia
cecrops Fabricius, 1793 )
378 beon (Stoll), 1782
379 cecrops (Fabricius ), 1793
poeas ( Hiibner ), “1806” [1811]
ab. gottschalki (Clark & Clark), 1938
TMOLUS Hiibner, “1816” [1819]
Type: Papilio echion Linnaeus, 1767
380 echiolus (Draudt), 1920

echion Auctorum (nec Linnaeus, 1767)
381 azia (Hewitson), 1873

nipona Auctorum (nec Hewitson, 1877)
CALLOPHRYS (INCISALIA) Scudder, 1872
Type: Licus niphon Hiibner, [1819-] 1823 [1823]

VOLUME 24, NUMBER 1 ol

382

383

384

385

386

387
388

389

polios Cook & Watson, 1907
ab. davisi Watson & Comstock, 1920
irus (Godart), “1819” [1824]
a i. irus (Godart), “1819” [1824]
arsace (Boisduval & Le Conte ), [1833]
ab. baltaeta Scudder, 1889
balteata dos Passos, 1964 (lapsus calami)
b i. hadros Cook & Watson, 1909
henrici (Grote & Robinson ), 1867
a h. henrici (Grote & Robinson ), 1867
b_h. turneri Clench, 1943
c h. margaretae dos Passos, 1943
d_h. solatus Cook & Watson, 1909
fotis (Strecker), “1877” [1878]
f. mossii (H. Edwards ), 1881
f. schryveri Cross, 1937
_f. bayensis R. M. Brown, 1969
f. doudoroffi dos Passos, 1946
f. windi Clench, 1943
f. fotis (Strecker ), “1877” [1878]
augustinus ( Westwood ), “1846-52” [1852]
augustus (Kirby nec Fabricius, 1793), 1837
. augustinus ( Westwood ), “1846-52” [1852]
. helenae dos Passos, 1943
. croesioides Scudder, 1876
. iroides ( Boisduval), 1852
ab. immaculata (Cockle ), 1910
e a. annetteae dos Passos, 1943
lanoraieensis Sheppard, 1934
niphon (Hitbner), [1819—] 1823 [1823]
nipha ( Morris ), 1860 (lapsus calami)
a n. niphon ( Htbner), [1819-] 1823 [1823]
plautus (Scudder ), 1876
b_ n. elarki Freeman, 1938
eryphon (Boisduval), 1852
a e. eryphon ( Boisduval), 1852
b_ e. sheltonensis Chermock & Frechin, “1948” [1949]

moiAo»o ®

so 2 » &

CALLOPHRYS (SANDIA) Clench & Ehrlich, 1960
Type: Callophrys (Sandia) mcfarlandi Ehrlich & Clench, 1960

390

macfarlandi Ehrlich & Clench, 1960 (emendatio )
mcefarlandi Ehrlich & Clench, 1960

32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CALLOPHRYS (XAMIA) Clench, [1961]
Type: Thecla xami Reakirt, “1866” [1867]
391 xami (Reakirt), “1866” [1867]

blenina (Hewitson ), 1868

CALLOPHRYS (MITOURA) Scudder, 1872
Type: Thecla smilacis Boisduval & Le Conte, [1833] (= Papilio damon
Stoll, 1782 nec [Denis & Schiffermiiller], 1775 = Lycus gryneus
Hubner, “1816” [1819] )
392 loki (Skinner ), 1907
393 hesseli Rawson & Ziegler, 1950
394 gryneus (Hibner), “1816” [1819]
a g. gryneus (Hiibner ), “1816” [1819] (gen. vern. )
damon (Stoll nec [Denis & Schiffermiiller], 1775), 1782
demon (Skinner ), 1897 (lapsus calami )
damastus (Godart ), “1819” [1824]
auburniana ( Harris), 1862 ( partim)
ab. octoscripta Buchholz, 1951
gen. aest. smilacis ( Boisduval & Le Conte ), [1833]
auburniana ( Harris ), 1862 ( partim )
patersonia ( Brehme ), 1907
bg. sweadneri Chermock, “1944” [1945]
c g. castalis (Edwards), “1870-1” (1871)
discoidalis (Skinner ), 1897
form brehmei Barnes & Benjamin, 1923
395 spinetorum (Hewitson), 1867
ninus (Edwards ), “1870-1” (1871)
cuyamaca (Wright ), 1922
396 siva (Edwards), “1874-6” (1874)
a 8. siva (Edwards), “1874-6” (1874)
rhodope (Godman & Salvin), (1887)
bs. juniperaria J. A. Comstock, 1925
c s. mansfieldi Tilden, 1951
397 nelsoni ( Boisduval), 1869
a n. nelsoni ( Boisduval), 1869
ab. exoleta (H. Edwards ), 1881
b n. muiri (H. Edwards), 1881
398 johnsoni (Skinner), 1904

CALLOPHRYS (CALLOPHRYS) Billberg, 1820
Type: Papilio rubi Linnaeus, 1758

VOLUME 24, NUMBER 1 33

Lycus Hubner, “1816” [1819] (nec Lycus Fabricius, 1787)
Licus Hubner [1819-] 1823 [1823]
399 affinis (Edwards ), 1862
a a. washingtonia Clench, 1944
b a. affinis (Edwards ), 1862
400 sheridanii (Carpenter ), 1877 (emendatio )
sheridonii (Carpenter ), 1877 (lapsus calami)
_ a sheridanii (Carpenter ), 1877 (emendatio )
b s. neoperplexa (Barnes & Benjamin ), 1923
c s.newcomeri Clench, 1963
401 dumetorum (Boisduval ), 1852
a d.dumetorum (Boisduval), 1852
perplexa Barnes & Benjamin, 1923
402 apama (Edwards), 1882
a a. apama (Edwards), 1882
b a. homoperplexa Barnes & Benjamin, 1923
403 ecomstocki Henne, 1940
404 lemberti Tilden, 1963
405 viridis (Edwards), 1862

CALLOPHRYS (CYANOPHRYS) Clench, [1961]
Type: Strymon agricola Butler & Druce, 1872
406 miserabilis Clench, 1946
pastor Auctorum (nec Butler & Druce, 1872)
407 goodsoni (Clench), 1946
facuna Auctorum (nec Hewitson, 1877 )

ATLIDES Hubner, “1816” [1819]
Type: Papilio halesus Cramer, “1779” [1777]
Brangas Hubner, “1816” [1819]
408 halesus (Cramer), “1779” [1777]
a_h. halesus (Cramer ), “1779” [1777]
dolichos (Hubner ), 1823
juanita (Scudder ), 1868
b_h. eoreorani Clench, 1942
form estesi Clench, 1942

DOLYMORPHA Holland, 1931
Type: Thecla jada Hewitson, “1862” [1867 |
409 jada ( Hewitson ), “1862” [1867]
EURISTRYMON Clench, [1961]
Type: Thecla favonius Smith, 1797

34

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

410 polingi (Barnes & Benjamin ), 1926
411 favonius (Smith), 1797
412 ontario (Edwards), “1868-9” (1868 )

o. ontario (Edwards), “1868-9” (1868 )
o. violae (Stallings & Turner ), 1947
o. autolyeus (Edwards ), “1870-1” (1871)
o. ilavia (Beutenmiiller ), 1899

mirabelle (Barnes ), 1900

(ely @ top

HYPOSTRYMON Clench, [1961]

Type: Thecla critola Hewitson, 1874

413 eritola (Hewitson ), 1874
PANTHIADES Hibner, “1816” [1819]

Type: Papilio pelion Cramer, “1779” [1775]

Parrhasius Hubner, “1816” [1819]
Eupsyche Scudder, 1876

414 m-album (Boisduval & Le Conte), [June 1833]

8 psyche (Boisduval & Le Conte) [June 1833]

STRYMON Hiibner, 1818 (opinion 541, name 1332)
Type: Strymon melinus Hubner, 1818 (opinion 541, name 1602)

415
*416

417

Bithys Hubner, 1818 ( opinion 541, name 1234 )
Bythis Geyer, [1827-31] (opinion 541, name 1236) (lapsus
calami )
Argus Gerhard (nec Bohadsch, 1761, nec Scopoli, 1763, nec
Scopoli, 1777, nec Boisduval, 1832), 1850
Callipareus Scudder, 1872
Callicista Grote, 1873
Uranotes Scudder, 1876
martialis ( Herrich-Schaffer ), 1864
acis ( Drury ), “1770” [1773]
mars ( Fabricius ), [1777]
a a. bartrami (Comstock & Huntington ), 1943
melinus Hubner, 1818 (opinion 541, name 1602)
a m. melinus Hubner, 1818
hyperici ( Boisduval & Le Conte), [27 July 1833]
ab. youngi Field, 1936
b m. humuili (Harris), 1841
ab. meinersi Gunder, 1927
c m. franki Field, 1938
d m. pudieca (H. Edwards), 1877

VoLUME 24, NUMBER | 35

e m. atrofasciata McDunnough, 1921
f m. setonia McDunnough, 1927
418 avalona (Wright), 1905
419 rufofusea (Hewitson ), “1862” (1877) (emendatio )
rufo-fusca ( Hewitson ), “1862” (1877)
420 cestri ( Reakirt), “1866” [1867]
421 yojoa (Reakirt), “1866” [July 1867]
*422, columella (Fabricius ), 1793
a ¢. istapa ( Reakirt), “1866” [1867]
b ec. modesta (Maynard), 1873
ocellifera (Grote ), 1873
493 bazochii (Godart), “1819” [1824]
thius Geyer, 1832
agra Hewitson, 1871
494 alea (Godman & Salvin), “1879-1901” [1887]
laceyi (Barnes & McDunnough), 1910
425 bebrycia ( Hewitson), 1868
buchholzi H. A. Freeman, 1950

ERORA Scudder, 1872
Type: Thecla laeta Edwards, 1862
496 laeta (Edwards), 1862
2 clothilde (Edwards ), “1863-4” (1863)
*427 quaderna ( Hewitson), 1868
a gq. sanfordi dos Passos, 1940

ELECTROSTRYMON Clench, [1961]
Type: Papilio endymion Fabricius, 1775
*428 endymion (Fabricius), 1775
a e. cyphara ( Hewitson), 1874

TAXONOMIC DISCUSSION
The following names, arranged in alphabetical order, have been
omitted from this revised synonymic catalogue because they are now
believed not to occur in the Nearctic region:

PLycaena erytalus Butler, “1869” [1870] (?lapsus calami for Tmolus
eurytulus Hiibner, 1819 )

Polyommatus hugon Godart, “1819” [1824]

Strymon eurytulus (Butler), “1869” [1870] is a South American species
of which erytalus auctorum is a misspelling and these names are ac-
cordingly omitted from the list.

36 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Strymon daraba (Hewitson ), “1861” [April 1867] is also a South American
species and dropped from the List.
Strymon pastor Butler & Druce, “1869-76” [1872]
Thecla beroea Hewitson, 1868
Thecla cybira Hewitson, “1862-78” [1874]
Thecla facuna Hewitson “1862” [1877]
Thecla nipona Hewitson, 1877
The following explanations will help answer questions concerning
placement of names, especially where the above list deviates from the
treatment of Clench, or where more recent information has been in-
corporated.

HARKENCLENUS

The International Commission on Zoological Nomenclature by opinion
541 suppressed among other names Chrysophanus Hubner, 1818, and
placed it on the Official Index of Rejected and Invalid Names in Zoology
for the purposes of the Law of Priority but not for those of the Law of
Homonymy. Consequently, a replacement name is in order. For that
purpose Harkenclenus has been chosen, being an arbitrary combination
of the first syllables of the name of my friend and colleague, Harry
Kendon Clench. The new name is masculine.

SATYRIUM

Satyrium edwardsii has from time to time been ascribed to Saunders
or Grote & Robinson. It was a manuscript name of Saunders first pub-
lished by Grote & Robinson (1867) in the synonymy of S. calanus. Since
that is no longer considered a valid publication (Code, Art. 16 [b] [ii]),
the name must be credited to Saunders who validly published it in 1869.

KUMAEUS

Clench removed Eumaeus from the Thecliti, and placed it in the
Strymoniti.

There are differences of opinion concerning the correct spelling of the
name of the type species of this genus. When first published, it was
written minijas, but when Hiibner later proposed this generic name and
included this species in it, he wrote the name minyas, and that spelling
has been generally accepted as a justified emendation. It seems better to
adhere to the present practice especially since Minyas was a Greek hero
and the letter “j” was not included in the classical Latin alphabet.

HETEROSMAITIA
Thecla spurina Hewitson, “1862” [1867] and T. zebina Hewitson
?
1862" [1567] were both named from single specimens taken in the

VOLUME 24, NUMBER 1 on

Amazon and Nicaragua respectively. One specimen of each has been
reported from southern Texas by Stallings & Turner and H. A. Freeman.
According to Clench in Ehrlich & Ehrlich ([1961]: 198), “Their true
identity remains uncertain.” The former according to Clench (in litt.)
belongs to the genus Heterosmaitia Clench, 1964, and it is probable that
the latter belongs to the same genus. He is unable to place them with
certainty at present.

CALLOPHRYS

Clench in Ehrlich & Ehrlich treats Callophrys viridis as a species in
his key but omits it from the text. It is here included as a valid species.
Further studies, on the biology and larval morphology, by G. A. Gorelick
(J. Lepid. Soc., in press) have tended to confirm this separation.

ATLIDES

The correct taxonomic standing of estesi presents a problem of nomen-
clature. The name was proposed as “Atlides halesus corcorani, form
estesi, new normal form” (Clench, 1942). From a reading of Clench’s
paper, it is clear that he intended estesi to represent the western popula-
tion of halesus. On the other hand, Gunder had proposed corcorani as a
transitional form (1934: 131). Transitional forms are generally considered
aberrations, but Clench by his action gave this one subspecific standing.
Accordingly, these names have been left as they stand in the synonymic
list, but the authorship of corcorani has been ascribed to Clench, 1942.

LITERATURE CITED

Ciencu, H. K., 1942. A new race of Atlides halesus Cramer from California
(Lepidoptera: Lycaenidae). Ent. News, 53: 219-221.

[1961]. Subfamily Lycaeninae. Blues, coppers, harvesters, and hairstreaks. In:
Ehrlich and Ehrlich. How to know the butterflies. Wm. C. Brown Company,
Dubuque, Iowa, [8] + 262 pp., 525 figs.

1963. A synopsis of the West Indian Lycaenidae with remarks on their zooge-
ography. J. Res. Lepid., 2: 247-270.

pos Passos, C. F., 1964. A synonymic list of the Nearctic Rhopalocera. Memoir
No. 1, The Lepidopterists’ Society, vi + 145 pp.

1969. <A revised synonymic catalogue with taxonomic notes on some Nearctic
Melitaeinae. J. Lepid. Soc., 23: 115-125.

Downey, J. C., [1961]. Tribe Plebejini. The blues. In: Ehrlich and Ehrlich.
How to know the butterflies. Wm. C. Brown Company, Dubuque, Iowa, [8] +
262 pp., 525 figs.

EuruicH, P. R., anp H. K. Ciencu, 1960. A new subgenus and species of
Callophrys (s.l.) from the southwestern United States (Lepidoptera: Lycaenidae).
Ent. News, 71: 137-141, pl. 1.

FREEMAN, H. A., 1950. A new species of hairstreak and new records for the
United States (Lepidoptera, Rhopalocera, Lycaenidae). Field and Lab., 18:
12-15.

Grote, A. R., AND C. T. Ropinson, 1867. Descriptions of American Lepidoptera.
—No. 2. Trans. Amer. Ent. Soc., 1: 171-192, pl. 4.

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Gunoer, J. D., 1934. Various new butterflies (Lepid., Rhopalocera). Can. Ent.,
66: 125-131.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE, 1959. Francis Hem-
ming, ed. Opinion 541. Suppression under the plenary powers of the generic
names Chrysophanus Hiibner, 1818, and Bithys Hiibner, 1818 (Class Insecta,
Order Lepidoptera) (Opinion supplementary to Opinion 165). Opinions and
declarations rendered by the International Commission on Zoological Nomencla-
ture, 20: 90-101.

STALLINGS, D. B., AND J. R. Turner, 1947. Texas lepidoptera (with description
of a new subspecies). Ent. News, 58: 36-41.

ON THE NATURE AND USE OF THE SUFFIX -ELLUS, -ELLA,
-ELLUM IN SPECIES-GROUP NAMES

GEORGE C. STEYSKAL

Systematic Entomology Laboratory, Entomology Research Division,
Agricultural Kesearch Service, U.S. Department of Agriculture?

The use of the suffix -ellus (-ella, -ellum) to form species-group names,
especially in Microlepidoptera in the form of -ella, has been common for a
long time, but fortunately it now seems to be waning. There is nothing to
be gained by adding this suffix to specific names, except a few letters to
the length of the name. Of the approximately 950 names cited in Hein-
rich’s revision of the American Phycitinae (1956), for example, more than
500 are formed with this suffix, starting with Tinea abietella Denis and
Schiffermueller, 1776,2 Tinea convolutella Hubner, 1796, Tinea decuriella
Hiibner, 1796, and Tinea elutella Hiibner, 1796, and continuing to Ambesa
columbiella McDunnough, 1935, and Epischnia vividella McDunnough
1935. None of the 89 new names proposed by Heinrich in this work are
formed in this wise.

Latin grammars deal with -ellus, -a, -um as follows:

a)... “diminutives (with endings for gender), forming nouns or adjectives, mean-
ing little or tender: as... puella, asellus, misellus.” (Allen and Greenough, 1872).

b) “There is a class of derivatives called diminutives, which express smallness:
lence also sometimes endearment, contempt, pity, or depreciation. Such are of the

' Mail address: c/o U.S. National Museum, Washington, D. C. 20560.

“lam indebted to R. W. Hodges for pointing out that the ending dates back at least to the 10th
edition of Linnaeus’ Systema Naturae, where the genus Phalaena is divided into 7 sections, in some
of which the specifie names regularly bore characteristic endings: Bombyces, Noctuae, Geometrae

and ata), Tortrices (-ana), Pyralides (-alis), Tineae (-ella), Alucitae (six species: mon-
tyla, didactyla, tridactyla, tetradactyla, pentadactyla, hexadactyla). These sections were dis-
hed a manner similar to that in which we now designate subgenera, e.g., Phalaena (Tortrix)
me nana, P ( ] inea) bella, P. (Tinea) euonymella. The Linnean sections now correspond roughly
nilies. The endings were used by later authors in various families. The endings -ana and -alis
\djective ‘nia and -ata are used with both nouns and adjectives; and -dactyla is the second

of compounds, which with the numerical 1st members form adjectives.

VOLUME 24, NUMBER | 39

same gender as their primitives. They are formed by adding 1. -ulus, -ula, -ulum . . .;

2. -culus, -cula, -culum ... Note 2.—A contraction is sometimes formed (especially
when the clipt stem of the primitive ends in 1, n, or r), and the termination is changed
into -ellus, -ella, or -ellum; . . . oculus, an eye, ocellus, a dear little eye; catulus, a

puppy, catellus, a little puppy; populus, the people, popellus, the rabble . . . Note 4.—
Adjectives also have diminutives: as miser, misellus.” (Chase, 1882).

c) “Diminutives, either nouns or adjectives, are formed from nouns or adjectives
with the endings . . . ellus, in any or all genders ... The gender of the primitive word
is generally retained in the diminutive. (Examples) castellum, tabella, libellus, puella,
flagellum.” (Jenks, 1911).

d) “From the stems of other nouns... El-lus, el-la, el-lum, . . . are used when the
stem of the primitive ends in a or 0, preceded by |, n, or r: ocellus ..., fabella .. .
Adjectives from adjectives .. . Note 1—The endings ellus and illus also occur as nouns:
novellus.” (Harkness, 1892).

e) “The diminutive ending -ellus is due to phonetic changes: as agellus (ager);
fabella (fabula) . . . Diminutive adjectives are formed like diminutive nouns, and
with a similar variety of meaning: pulchellus (from pulcher).’ (Burton, 1911).

Cooper (1895), in an extended work on word formation in popular
Latin, devotes 33 pages (pp. 164-195) to diminutives, pointing out that
they lost strictly diminutive meaning at an early date, that such weakening
of the meaning continued throughout the history of the language, that in
later periods a fondness for compound, 2nd and 3rd degree diminutives
became more evident, and that diminutive adjectives were far less numer-
ous than diminutive nouns. He cites many examples.

Stern (1966, p. 289) remarks, in regard to generic names in botany,
that “... the suffix -ella has now come to be regarded, particularly by
mycologists, as being simply a name-forming component to be attached to
any personal name or any generic name of either Latin or Greek origin,
usually without implication of smallness.” The same condition is true
with regard to generic names in entomology, and at least in the Micro-
lepidoptera the condition has been carried over into species-group names.
The following examples, all from Heinrich (1956), will show this.

A) From nouns: abietella (Abies), arizonella (Arizona), bahamasella (Bahamas),
bacerella (Baker), bakerella (Baker), cacabella (cacabus), castrella (castrum = Fort
[Wingate] ), cnabella (Knab), constitutionella (constitution = constitutio), crataegella
(Crataegus), exsulella (exsul), fasciella (fascia), gitonella (Greek geitén), glendella
(Glenwood [Springs] ), homoeosomella (Homoeosoma), illuviella (illuvies), titillella
(titillus ).

B) From adjectives: aeneella (aenea), abietivorella (abietivora), albescentella
(albescens ), albidiorella (albidior), angustellus (angustus), atrella (atra), australella
(australis ), cinerella (cinerea), glabrella ( glabra; better glabella), melanellus (melas,
melan-).

C) From ?: cinerella (? noun cinis, ciner-; ? error for cinereella), demotella (?
Greek noun démotés ); immorella; jocarella; obnupsella; obsipella (? verb obsipo ); op-
oredestella (? Greek nouns opora + edestés); plorella (? verb ploro); senesciella (?
verb senesco ), verecuntella (? adj. verecunda ).

With generic names ending in -ellus, -ella, -ellum, there is no trouble,

since these endings clearly show the gender. But with species-group

40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

names, it is necessary to know whether the name is noun or adjective
when it is transferred to a genus of different gender. We cannot simply say
that the names are either all nouns or all adjectives. It is not possible to
make good Latin words like pulchella, atrella, or angustellus, into anything
but adjectives, nor is it possible to make anything but nouns (invariable as
to gender) out of such words as cacabella, exsulella, castrella, bakerella,
arizonella, etc. With names that are obviously either nouns or adjectives
by original stated derivation, by Latin or Greek dictionaries, by derivation
from a generic name, etc., there is a clear choice, but with words such as
those cited in category “C” above, no source yields any usable data for the
choice between noun and adjective.

As an example of the problems that could be encountered, let us suppose
that 7 names recently proposed (Shaffer, 1968 ) were to be transferred to a
genus of different gender. These names are Anacostia tribulella, Arivaca
artella, A. linella, A. poohella, Homosassa incudella, H. platella, and Peoria
floridella. No derivations were given for these names, but personal com-
munication with the author brought out the information that floridella
was based upon the name of the State of Florida, that incudella was based
upon the Latin verb incudo (“because the male genitalia have a part re-
sembling an anvil”), and that the others are neologisms, or made-up
names. None of these names contravenes any of the International Rules
of Zoological Nomenclature; they are therefore validly formed and even
rather good names from the standpoints of simplicity, shortness, distinc-
tiveness, and ease of pronunciation for most people.

Many zoologists nowadays are simply coining names rather than going
through a rather laborious and time-consuming process of looking for
something apt and linguistically correct in a language with which they
lack familiarity. But that course, too, has its limits and dangers. One
might come up with the names alba and leo by shuffling a set of letter
cards, but the fact that those words have been good Latin words for many
centuries cannot be controverted. They now belong to all mankind and
cannot be said by any one person to mean anything other than what their
usage as recorded in lexicons has been all this time.

Therefore, of Shaffer's names, one must be considered an adjective
(artella, from the Latin adj. artus, -a, -um close, narrow, confined ) and 5
others are clearly nouns (floridella, from Florida rather than from the
Latin adj. floridus, -a, -um flowery; tribulella, from Latin noun tribulus:
incudella, from the Latin noun incis anvil, with gender change [the verb
incudo is from the same root]; linella, from the Latin noun linum flax,
thread, line, rope, with gender change; and platella, from either of the
Greek nouns platé blade, flat part of an object, or platos breadth, but not

VOLUME 24, NUMBER 1 Al

from the adj. platys, which would give platyella). This last name could
have been formed from the generic name Platus Motschulsky, 1844
(Coleoptera ), far-fetched of course, but that would also make platella a
noun.

The remaining name, poohella, is rather obviously made-up; at least I
can find no classical basis for it, nor does there seem to be a genus-name it
could have been based upon. Since it is a neologism and certainly not
derived from any Latin or Greek adjective, it is best considered as a noun,
the most basic part of language. In order to be an adjective, it would have
to have a meaning of adjectival nature. Any word, however, can be used as
a noun. Latin adjectives had the capability of usage as nouns, but as
species-group names in biological nomenclature what is by nature an ad-
jective can have nothing like Latin sentence structure to show that it is
being used as a noun. The names obsipella and plorella (see above, from
Heinrich, 1956), being traceable only to a verbal root, are also best con-
sidered as neologistic nouns.

As long as the International Rules and the Latin language are what they
are, indication of the derivation of newly formed names is highly to be rec-
ommended to save other workers much time and effort that might be
used much more profitably. The tracing of such a word as cacabella to the
obscure Latin word cacabus, found in only the most complete lexicons, or
the word incudella to the noun inciis, with its hidden root form incud-, is
certainly not a very useful occupation.

More concise statements in the International Rules regarding availabil-
ity and treatment of non-Latin specific names would also do much to
obviate growing confusion.

LITERATURE CITED

ALLEN, J. H., & J. R. Greenoucu, 1872. A Latin grammar for schools and colleges,
founded on comparative grammar. Ginn Bros., Boston, xv + 252 pp.
Burton, H. E., 1911. A Latin Grammar. Silver, Burdett and Co., N. Y., etc., 337

pp.

Case, T., 1882. A Latin Grammar. Eldredge and Bro., Philadelphia, viii + 313 pp.

Cooper, F. T., 1895. Word formation in the Roman Sermo Plebeius. Privately publ.,
N. Y. (thesis, Columbia College, N. Y.), xlvii + 329 pp.

Jenks, P. R., 1911. A manual of Latin word formation for secondary schools. D. C.
Heath and Co., Boston, etc., (v) + 81 pp.

Harkness, A., 1892. A Latin grammar for schools and colleges. Rev. std. ed., Amer.
Book Co., N. Y., etc., 430 pp.

HErIneicH, C., 1956. American moths of the subfamily Phycitinae. U. S. Nat. Mus.
Bull. 207: viii + 581 pp.

SHAFFER, J. C., 1968. A revision of the Peoriinae and Anerastiinae (auctorum) of
America north of Mexico (Lepidoptera: Pyralidae). U.S. Nat. Mus. Bull. 280:
vi + 124 pp.

STERN, W.T., 1966. Botanical Latin. Hafner Publ. Co., N. Y., xiv + 566 pp.

42 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

NEW MICHIGAN BUTTERFLY RECORDS

M. C. NIELSEN
3415 Overlea Drive, Lansing, Michigan

Since the death of Sherman Moore in 1956, and the publication of his
list of Michigan butterflies (1960), the writer has assumed the task of
maintaining a card-file index of all Michigan butterfly records, with the
goal of preparing a new annotated list. During this period, considerable
collecting has occurred throughout the state, particularly in the Upper
Peninsula and on Isle Royale. This period has seen a definite increase in
the number of resident lepidopterists and young insect collectors, especially
among the 4-H group. Therefore, it seems worthwhile to publish certain
new significant records at this time. Hopefully, this addendum may en-
courage lepidopterists to search diligently in the state for additional
species to further the knowledge of the distribution and habits of Michi-
gan’s butterfly fauna.

Field trips to Isle Royale by David Bixler, Edward Voss and Ron Will-
son have added immeasurably to our meager knowledge of the island’s
butterfly fauna, which incidentally appears to be similar to that of northern
Minnesota and northern Ontario. Collecting by Virgil Warczynski of Bay
City is responsible for many new county records, particularly in the
Saginaw Bay area. Brief collecting forays by John Masters of Minnesota
into the western portion of the Upper Peninsula have produced interesting
new records; John Newman and the writer have done extensive collecting
in the numerous sphagnum-heath bogs in the eastern Upper Peninsula
since the publication of Moore’s list. Additional Upper Peninsula records
were found in the insect collections at Northern Michigan University in
Marquette and Michigan Technological University in Houghton.

As a result of these additional records, we can now add over 300 new
county records, including 7 new species (noted with an asterisk) to our
list. Only those new county records which significantly extend a species’
range, or relate to a scarce or extremely local species are treated in this
supplement. Also, there is recognition of certain subspecies heretofore
omitted by Moore. With the addition of these new state records, our
Michigan butterfly fauna now constitutes 147 recognized species. Hope-
fully, further collecting on Isle Royale, in the Upper Peninsula, and in the
southwestern Lower Peninsula should eventually bring the state list to at
least 155 species. Knowledge of any new state or county records not
previously published from Michigan would be appreciated by the writer.

‘n presenting the following records, the nomenclature and arrangement

VOLUME 24, NUMBER 1 43

of Klots (1951) has been adopted in order to maintain conformity to
Moore's list.

SATYRIDAE

Euptychia mitchellii (French). Branch, Lenawee Counties, June 30, July
11. These two locations were previously unrecorded (McAlpine, Hubbell,
Pliske, 1960). Additional data is still needed to delineate its exact range in
Michigan, especially the northern limits.

Oeneis macounii (Edwards ). Isle Royale, June 27, 1964. A fresh pair was
collected by Ron Willson in Section 22, T66N, R34W; other specimens
were observed in this section on Greenstone Ridge. The specimens are
deposited in the Michigan State University collection at East Lansing. This
is the first record of the species from the main island since its discovery on
one of Isle Royale’s satellite islands by R. N. Rysgaard (Newcomb, 1941).
It would indicate that macounii is a resident on Isle Royale and should be
searched for elsewhere on the island, as well as in suitable habitats in
the Upper Peninsula. The butterfly has a tendency to resemble Limenitis
archippus (Cramer) in flight (Fletcher, 1888), and inhabits open jack
pine forests (Masters, Sorensen, Conway, 1967).

Oeneis jutta ascerta Masters & Sorensen. Chippewa, Mackinac Counties,
May 29, June 15. Several specimens were collected and observed in black
spruce-tamarack bogs ( Nielsen, 1965). This recently described subspecies
(Masters, Sorensen, 1968) should also occur on Isle Royale and in many
acid bogs throughout the Upper Peninsula.

* Erebia discoidalis (Kirby). Baraga County, May 31, 1968. One torn
male collected by J. H. Masters in a black spruce bog around 5:00 PM. Ac-
cording to correspondence with Masters, discoidalis apparently flies early
in the morning and late in the afternoon in most northern bogs, a habit that
could account for its scarcity to date. This alpine should occur in many
bogs across the Upper Peninsula, as it has been previously collected in
Minnesota (Huber, 1965) and Wisconsin (Elder, 1961). The Michigan
specimen has been deposited in the MSU collection.

NYMPHALIDAE

* Boloria eunomia dawsoni (Barnes & McDunnough). Chippewa,
Mackinac Counties, June 13-23. The writer has taken additional speci-
mens and located two new colonies in Chippewa County since its dis-
covery (Nielsen, 1965).

* Boloria freija (Thunberg). Chippewa, Ontonagon Counties, May 27,
29. The first record was that of a worn male collected by John Newman in
Section 9, T49N, R7W in 1965, on a sandy ridge extending across the huge

44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bog located about seven miles northwest of Paradise (Nielsen, 1965).
More recently, it was taken by John Masters in the eastern edge of the
Porcupine Mountains State Park.

Boloria frigga (Thunberg). Schoolcraft County, May 30-June 1. Addi-
tional specimens have been collected which indicate that the species may
prefer a wet, sedgy bog habitat, with willow and dwarf birch, to the true
sphagnum-heath bog previously reported (Hubbell, 1957). The writer
has collected it in a habitat similar to that reported by Brown (1954) for
frigga sagata (Barnes & Benjamin ) in the Rocky Mountains of Colorado.

* Polygonia zephyrus (Edwards). Schoolcraft County, May 29, 1960.
One torn male of this western species was collected in an acid bog by the
writer (Section 14, T42N, RI6W). The specimen was determined by
Cyril F. dos Passos and is now in the writer's collection. It would appear
this record represents a stray specimen which probably overwintered as
suggested by its condition. Undoubtedly, bait could be used to attract
additional material, including other scarce boreal species as P. gracilis
(Grote & Robinson ) and satyrus marsyas (Edwards ).

Anaea andria Scudder. Van Buren County, July 17. This record represents
the second location in the state. One specimen was collected by William
Bouton at Breedsville. Collectors should look for it in other southern
counties, especially at bait.

LYCAENIDAE

* Strymon m-album ( Boisduval & LeConte ). Muskegon County, August
12, 1964. One fresh male, Michigan’s first, was collected by John Taggart
in Section 13, TLIN, R18W, along a sandy road adjacent to scrub oak. The
specimen is in the writer's collection. This hairstreak may occasionally
occur in our southern scrub oak locations; although it is doubtful that it
breeds in the state. According to Clench (1961), the species is triple-
brooded in Pennsylvania.

Strymon melinus humuli (Harris). Baraga, Chippewa Counties, May 17,
June 21. These are the first records of this hairstreak collected from the
Upper Peninsula and they constitute a substantial northern extension. Two
orn specimens were taken by the writer near Strongs in an open jack pine
ing with Incisalia niphon clarki Freeman and Hesperia sassacus

Incisalia henrici (Grote & Robinson). Schoolcraft County, June 1. This is
the fi ‘ this elfin in the Upper Peninsula. John Newman col-

lected on imen along a sandy trail separating a jack pine ridge

VOLUME 24, NUMBER 1] 45

and an acid bog; Vaccinium sp., one of its known foodplants ( Klots, 1951 ),
was abundant in the area.

Incisalia eryphon (Boisduval). Chippewa, Luce, Marquette Counties,
May 16-22. The determination of these specimens has been confirmed by
Harry Clench (Nielsen, 1966) as representing the western pine elfin. The
Mackinac County specimens, referred to by Moore (1960), have since
been examined by the writer and were found to represent eryphon and not
niphon clarki as determined by Klots. There is good reason to believe that
eryphon will be found in pine areas throughout the Upper Peninsula, and
that it has previously been overlooked or confused with niphon clarki.

PAPILIONIDAE

Papilio glaucus canadensis Rothschild & Jordan. Keweenaw County, June
26. Five specimens taken by Ron Hodges at Copper Harbor were deter-
mined by Dr. Lincoln P. Brower. This is the first recognition of this sub-
species from Michigan; Moore (1960) did not distinguish this from the
typical form. No doubt the populations on Isle Royale and throughout the
Upper Peninsula and extending south into the Lower Peninsula represents
canadensis. Specimens collected from Otsego County are indistinguish-
able from those taken farther north. Further collections of long series
should be made to determine the southern limits of canadensis.

PIERIDAE

* Euchloe ausonides Lucas. Isle Royale, June 17-29. The species was
first collected by Ron Willson in 1964 near Rock Harbor Lodge at the
northeastern end of the island. Additional specimens were taken on Mt.
Ojibway by David Bixler, and again at Rock Harbor by Ed Voss. Corre-
spondence from Voss indicated that ausonides was associated with Arabis.
Willson’s specimen was determined by Dr. A. B. Klots, but was undeter-
minable as to subspecies. The species has also been collected in northern
Minnesota and in the vicinity of Port Arthur, Ontario.

Pieris virginiensis Edwards. Arenac, Otsego Counties, May 5, 26. Several
specimens were collected by Virgil Warczynski in hardwoods, thus extend-
ing the known range of this pierid to the southeast. Collectors should
look for this butterfly in beech-hard maple woodlots in early spring before
full leaf development.

HESPERIDAE

Pyrgus centaureae wyandot (Edwards ). Montcalm County, May 8, 18. The
writer has taken this skipper feeding on blueberry blossoms in scrub oak

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

openings in Section 7, T12N, R1OW. This represents the most southern
collection site in the state.

Erynnis baptisiae (Forbes). Monroe County, June 5. This skipper was
collected by the writer in Section 24, T7S, R6E, in close proximity to wild
indigo in scrub oak opening; subsequently, adults were reared from eggs
and larvae found on wild indigo plants in this same location. The species
is easily confused with our other Erynnis species.

Oarisma powesheik (Parker). Kent County, July 4, 13. Several specimens
have been collected in recent years by Wilbur McAlpine, V. Warczynski
and the writer along the marsh edge of Button and Lamberton Lakes in
the north section of Grand Rapids. Somehow, this small and restricted
colony has continued to flourish since first reported by Wolcott (1893)
despite the suburban encroachment. The writer found the species to be
rather fond of black-eyed susan flowers growing near the marsh perimeter.
Companion species taken in this habitat included Calephelis muticum
McAlpine and Lycaena dorcas Kirby. It would appear this species can be
easily overlooked in this type of habitat.

Adopaea lineola (Ochsenheimer). Cheboygan, Crawford, Houghton,
Oscoda, Otsego, Roscommon Counties, June 19-July 22. Several specimens
were collected by the writer in the northern Lower Peninsula during the
1960-67 period, and by Ted Ellis, Jr. in Houghton County; these represent
significant northern extensions in range for this skipper. Undoubtedly, this
skipper will spread to all counties of the state within the next several years.
The writer recently collected a long series in two northern Ontario loca-
tions ( Nielsen, 1966), indicating that the European Skipper is still extend-
ing its range in North America.

* Lerodea eufala (Edwards). Ontonagon County, August 25, 1959. One
torn female was collected by the late Dr. R. R. Dreisbach. Determination
of this skipper was confirmed by Dr. A. W. Lindsey; the specimen is in
the writer's collection. The skipper may be extending its range northward

into the western end of the Upper Peninsula and should be searched for in
upland grassy areas,

LITERATURE CITED

Brown, F’. Martin, 1954. Some Notes on Boloria in Central Colorado ( Nymphal-
idae). Lepid. News, 8: 64-66.

CLENC H, H. ] 961 Panthiades m-album (Lycaenidae): Remarks on Its Early

Stages and on Its Occurrence in Pennsylvania. Jour. Lepid. Soc., 15: 226=933)

“LDER, W. H 196] ‘arly

: | , tecords of Erebia discoidalis (Satyridae) in Wisconsin.
Jour. Lepid. Soc 95-96.

FLETCHER, J., 188! ‘rip to Nepigon. Ann. Report Ent. Soc, Ontario, 19: 74-88.

VOLUME 24, NUMBER | 47

Huser, R. L., 1965. Probable Second U. S. Record for Erebia discoidalis. Jour.
Lepid. Soc., 19: 76.

Hvusse.t, S.P., 1957. Boloria frigga (Nymphalidae) in Michigan. Lepid. News, 11:
37-38.

Kxiots, A. B., 1951. A Field Guide to the Butterflies of North America, East of the
Great Plains. Houghton Mifflin Co., Boston, 349 pp.

Masters, J. H., and J. T. SorENSEN & P. J. Conway, 1967. Observations on Oeneis
macounii (Satyridae) in Manitoba and Minnesota. Jour. Lepid. Soc., 21: 258—
260.

Masters, J. H., and J. T. SorENsEN, 1968. A New Subspecies of Oeneis jutta (Lep-
idoptera: Satyridae). Ent. News, 79: 80-85.

McA.PInE, W. S., and S. P. HuspELL & T. E. PiiskE, 1960. The Distribution, Habits,
and Life History of Euptychia mitchellii (Satyridae). Jour. Lepid. Soc., 14: 209-
226.

Moores, S., 1960. A Revised Annotated List of the Butterflies of Michigan. Occ.
Papers Mus. Zool., Univ. Mich., 617, 39 pp.

Newcomes, W. W. 1941. Note on the Occurrence of Oeneis macounii. Bull.
Brooklyn Ent. Soc., 36: 56.

NiELSEN, M. C., 1965. Discovery and Observations of Boloria eunomia (Nymphali-
dae) in Michigan. Jour. Lepid. Soc., 18: 233-237.

1966. Occurrence of Callophrys eryphon (Lycaenidae) in Michigan. Jour.
Lepid. Soc., 20: 41-42.

1966. New Canadian Records for Thymelicus lineola (Hesperiidae).
Jour. Lepid. Soc., 20: 243-244.

Wotcortt, R. H., 1893. The Butterflies of Grand Rapids, Michigan, Canad. Ent.,
25: 98-107.

HEAD MEASUREMENTS AND WEIGHTS OF THE
BEAN LEAF ROLLER, URBANUS PROTEUS (HESPERIIDAE )!

GERALD L. GREENE
Central Florida Experiment Station, Sanford, Florida 32771

The bean leaf roller, Urbanus proteus (Linn.), is common on beans and
other legumes in Florida during the fall of the year. Head widths and
weights of newly molted larvae were determined to learn if larval instars
could be accurately distinguished when making field observations.

Larvae were reared on Harvester variety snap bean leaves held in an
insect rearing room with the temperature about 85°F. Larvae were kept
individually on bean leaflets in round plastic dishes 2 cm deep by 10 cm in
diameter with moistened filter paper to maintain leaf turgidity. Fresh
leaves were supplied after each molt. Eggs were measured within five
hours after oviposition and larval head widths were recorded soon after
molting when the head became black. Pupal heads were measured two

1 Florida Agricultural Experiment Stations Journal Series No. 3241.

48 JoURNAL OF THE LEPIDOPTERISTS’ SOCIETY

6 HEAD WIDTH
5) oll

S) a7 a a
ae
=

Soff
= 3 | cs
=>
= 2 | 1.8
oll
/ I 0.6 |
- § aes
EGG: 3 4 5 PUPA ADULT

INSTAR

Fig. 1. Comparison of egg and head widths of the bean leaf roller, Urbanus proteus
(Linn.). Bars and numbers represent means of 50 or more individuals and the lines
connect the low and high measurement for each life stage.

days after pupation. All measurements were made on living specimens
except for adults, which were killed in a cyanide jar following emergence
and measured within one hour after death. Head measurements were
made using a dissecting microscope with an ocular micrometer.

Data on the means and ranges of head widths (Fig. 1) demonstrate
distinct differences between instars with none of the extreme measure-
ments overlapping from instar to instar. The validity of field determina-
tion of instars based on head size is substantiated by these data. The results
of these measurements agree with those reported by Quaintance (1898).
His measurements, made on preserved specimens, were as follows: eggs
(0.966 mm, Ist instar 0.566 mm, 2nd instar 0.866 mm, 3rd instar 2 mm, 4th
instar 3 mm, and 5th instar 4.5 mm. Quaintance did not report the ex-
tremes or variation within each stage.

Heads of newly eclosed larvae filled approximately % of the egg interior.
The 5th instar, pupal and adult head widths were similar (Fig. 1). When
observing the changes during these stages it appeared that pupal heads
would be wider than 5th instar larvae by a larger amount than was mea-
sured because the 5th instar head capsules split during pupation. The
adult emerges from the pupal case without splitting the head covering,
therefore its head cannot be wider than the pupal head. The extreme mea-

VOLUME 24, NUMBER 1 49

TABLE 1. Milligrams weight of eight life stages of the bean leaf roller, Urbanus
proteus (Linn.) taken at the beginning of each stage.

Egg? Ist? 2nd 3rd 4th 5th Pupa Adult?

Low individual — = ] 5 29 ILS 367 100
Mean 0.46 0.41 DM Ils BASS) IESG AoW il alg eo)
High individual — — 35 14 47 265 610 352
Weight change

ratio‘® — 0.9 5.5 5.0 3.5 5.5 1.9 (0.4
No. of individuals

weighed 170 120 126 WG 96 90 85 25

a Weighed in groups of 10 per weighing because the balance used was only sensitive to tenths
of a mg.

b Adults were killed with cyanide and weighed within a few minutes after emergence from the
pupa. All other weights were made of living individuals.

¢ The figures are derived by dividing the weight of the indicated stage by the weight of the
previous stage.

surements for the adult were much closer than they were for the 5th instar
or pupa.

Comparing head measurements by dividing the head width of one
instar into that of the preceding instar, ratios of .55, .61, .49, .79 and a mean
of .61 were found. Dyar (1890) found ratios of .58 to .73 for 14 species
having five larval stages, and the bean leaf roller falls within his range. A
lower numerical ratio was expected in an insect with a large head such as a
Hesperiidae, but this was not apparent with the bean leaf roller.

Dyar (1890) mentions the smaller size of head capsules from larvae
which died after molting. Several bean leaf roller larvae which died were
compared to the average of each instar. The first four instars which died
had similar measurements to live specimens, but the dead 5th instar heads
were much smaller than the average, and two which had not grown during
the 4th instar had heads smaller than the previous instar. Measurements
of live and preserved head capsules were very similar, even for 5th instar
heads which had split during pupation.

In conjunction with head measurements larvae were weighed to record
the weight increase from molt to molt. These weights were made just
after molting rather than before molting to allow for uniform weighing.
Weights just before molting would have been nearly impossible to make
due to the difficulty of knowing when the larvae had completed their
feeding. By weighing soon after molting, uniform weights were made for
individuals of each instar and feeding or molting was not interrupted.
Larvae had not begun to feed after molting when they were weighed.

The weights are given in Table 1 for the eight life stages. All weights

50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 2. Conversion of eaten leaf weight to bean leaf roller (BLR), Urbanus
proteus (Linn.), weight for five life stages.

Ege i 2 3 4 5 eapae
Leaf eaten (mg) — 11 28 110 567 3343 —
BLR weight (mg) 0.46 0.41 8) nie) 38.6 — DIS 2
BLR gain (mg)' — 1.9 8.9 27.2 186.6 SaiS2es —
Conversion ratio” — 0.17 0.32 0.24 0.33 0.06 —

1 This figure derived by subtracting weight of one instar from the weight of next instar. The
weight lost during molting is not included because weights were made after molting but before
feeding resumed.
mg BLR gain

a ; :
Bean : Bry oe ;
Conversion ratio mg Leaf eaten

are of live specimens, except for adults, which were killed just before
weighing. The ratio of gain was largest from Ist to 2nd and 4th to 5th
instar and smallest from 5th instar to pupa. The 5th instar gained con-
siderable weight, but lost much of the gain at pupation after which the
weights were taken. The small loss of weight from egg to newly eclosed
larva, only 10%, was a surprise, since newly eclosed larvae do not eat any
of the egg shell and less of fluids during eclosion was noted.

The conversion ratio of leaf weight to body weight of each larva was
calculated by dividing the weight of leaf tissue eaten by the weight in-
crease during each instar (Table 2). The efficiency of the lst and 5th
instar to convert leaf weight into body weight was low, whereas the ef-
ficiency of the other 3 instars was similar. The figure of 0.3 is close to the
conversion ratios reported by Taylor and Bardner (1968) for the diamond
back moth on a dry weight basis. The low conversion ratio for the 1st
instar was unexpected since larval weight increased 5.5 times over the
newly hatched larvae. After 5th instar feeding the transformation into
the pupa results in considerable weight loss as indicated by the 0.06 figure
in Table 1.

Przibram and Megusar (1912) concluded that the weight of each instar
should double that of the preceding instar and Bodenheimer (1932) added
latent divisions for holometabolic insects. Instar weights obtained for the
bean leaf roller agree closely with those reported by Przibram and
Megusar when the latent division calculations of Bodenheimer are used.

CONCLUSIONS
Measurements of bean leaf roller heads demonstrated that larval instars
can be accurately distinguished. The head widths did not overlap from

one instar to the next and the standard deviation was 0.4 mm for the 5th

VoLUME 24, NUMBER | bt

instar and pupae, 0.1 for adults and less than 0.1 for the other life stages.
Head capsules which had been shed during molting were similar in width
to live specimens.

Larval weight increased approximately 5 fold during each of the 2nd,
3rd, and 4th instar periods. The 5th instar increased in size and weight but
lost much of the increase during pupation. Adults weighed 0.4 times as
much as did the pupae, and Ist instar larvae averaged 0.9 times as much
as the eggs. A greater proportion of weight was lost during the transforma-
tion from 5th instar to adult than during eclosion.

LITERATURE CITED

BODENHEIMER, F. S. 1932. The progression factor in insect growth. Quart. Rev.
Biol. 8: 92-95.

Dyar, H. G. 1890. The number of moults of lepidopterous larvae. Psyche 5: 420-
422.

PrzirRaM, H., and F. Mecusar. 1912. Wachstumsmessungen an Sphodromantis
bioculata Burm. I. Lange und Masse. Arch. Entw. Mech. d. Organismen 34:
680-741.

Taytor, W. E., and R. Barpner. 1968. Leaf injury and food consumption by
larvae of Phaedon cochleariae (Coleoptera; Chrysomelidae) and Plutella maculi-
pennis (Lepidoptera, Plutellidae) feeding on turnip and radish. Entomol. Exp. &
Appl. 11: 177-184.

QuAINTANCE, A. L. 1898. Three injurious insects. Bean leaf-roller. Corn delphax.
Canna leaf-roller. Fla. Agr. Exp. Sta. Bull. 45: 49-62.

WHATS YOUR COLLECTION WORTH

CHARLES V. COVELL, JR.

University of Louisville, Kentucky

In discussing the value of a collection of Lepidoptera we must first
define “value” by some criterion. We can first consider its monetary
value: the amount spent on materials, storage equipment, library, and the
procurement of specimens; and also the fair market value if it were to be
sold to a dealer, private collector, or institution. Next, there is the senti-
mental value to the collector. Most collectors probably value their collec-
tions far beyond a fair market value simply because of all the hours of
sweating, searching, panting, itching, squinting, cursing, and joyful
whooping that accompany the perfect avocation. Finally, we shall discuss
the scientific value: what information useful in taxonomic and faunistic
research is intrinsic in the collection? The scientific value can sometimes
be related to monetary value; but, too often, institutional collections
cannot afford to purchase highly desirable collections, and must rely upon
donated material for research purposes.

52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The monetary value of a collection depends upon several factors, most
important of which is the quality of the material. Large collections, con-
taining showy and unusual exotic species, should command a good price
among the few dealers who buy and sell Lepidoptera. Material from
poorly-collected areas, unusual forms, type specimens, and unusual, well-
labeled perfect butterflies and moths will be in much more demand than a
small local collection of common species. There can be no fixed price for
specimens because of the variables of quality, rarity, and attractiveness as
well as supply and demand; but one can compare his material with that
offered at various prices by dealers to get some estimate of value, whether
for pricing his collection, or for evaluation for insurance purposes. Like-
wise, books and equipment can be evaluated by checking dealer price
lists.

One can realize monetary benefits by donating his collection to a major
museum or university collection. He can include the value of donated ma-
terial in the “miscellaneous contributions” section of Itemized Deductions
on his income tax returns. In checking with the Internal Revenue Service,
I found that IRS normally accepts the value placed on such donations by
an official of the recipient institution who will prepare a statement of
value for the donor. From experience and also in checking with an official
of one of the nation’s largest museums, I know that $.15 per specimen is an
acceptable average value for insects. I would think that one could justify
a slightly higher amount for spread Lepidoptera, because of the time put
into setting. Also, rare or otherwise extraordinary specimens can be given
much higher individual evaluations.

With respect to sentimental value, one cannot place a price tag on the
aesthetic enjoyment, friendships, and personal satisfaction of building a
fine collection. This intangible value is extended to others when they see
your specimens, hear you give a talk, or become stimulated to begin or
recommence collecting themselves. Although many collectors are satisfied
to keep their collections to themselves, I feel certain that those who share
their knowledge and experience reap much greater rewards. The pleasure
of excited responses from the uninitiated as well as fervent discussion with
fellow aficianados is its own reward.

The scientific value of a collection depends first upon the completeness
and accuracy of the data on pin labels or papers. Again, the rarity of
species, localities represented, and amount of type material are factors
important in determining what potential information the collection holds.
Condition of specimens is important, too; but it is not nearly as important
to the scientist as the dependability of the data furnished by the collector.

Unfortunately, there are many collectors who do not know that a speci-

VOLUME 24, NUMBER | 53

men without locality and other collection data is worthless to the re-
searcher. Others are careless and get dates and localities confused, or
deliberately mislabel specimens to enhance their prestige or line their
pockets. The notorious Chokoloskee, Florida, material is a good example
of the latter. Although some collectors will always be content to merely
place showy specimens in Riker mounts for their walls, I feel most would
prefer to prepare a collection that is scientifically as well as aesthetically
valuable. And the knowledge that scientific usefulness will also enhance
monetary value should act as an additional incentive to prepare accurate
labels and keep a field notebook.

Even small local collections can have significant value to science when
properly prepared. In many states faunistic knowledge is extremely
limited. With more and more land coming under the bulldozer, it is im-
portant that as much faunistic work on Lepidoptera as well as other biota
be completed as soon as possible. Since professional biologists often have
little time for collecting, the role of the amateur in contributing informa-
tion for taxonomic and faunistic studies is increasingly important.

Sadly enough, many fine collections are lost to science because the
collector failed to donate his collection while alive, or arrange for its
proper disposal in his will. One important North Carolina collection faded
away in display cases in the hallway of a state building; others end up
nourishing dermestids in high school labs or family attics. In order to
prevent such a loss of your collection, I offer these guidelines:

1. Keep your collection in good order. This includes proper labeling
(avoiding “coding” of specimens, and poor quality paper and ink), and
storage in air-tight, regularly-fumigated containers away from excessive
light and moisture.

2. Donate your collection while you are alive. Tf you are no longer
actively working on your collection, give it to an institution (or actually in-
corporate it in person) so that accession will be most efficiently effected.
In so doing, you may reap a tax benefit, recover storage equipment to sell
or give to a promising beginner, and feel secure that your specimens are in
responsible hands of your own choosing.

3. Spell it out in your will. If you do not donate your collection now, be
sure that you have provided for it according to the laws of your state. Ex-
plicit additional directions can be left in writing to aid the recipient in
understanding any confusing aspect of the collection; or, better still,
arrangements can be discussed with the recipient in advance.

4. Choose an appropriate recipient. Although you may prefer to give
your collection to a colleague, it is usually better to donate or bequeath it
to a large museum or university collection where proper care and use of

54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

. . . (<4 > ¢
your material are assured. Beware of institutions where “perpetual care” is
uncertain.

There are certainly other helpful ideas to aid you in evaluating and
utilizing your collection for more than the mere pleasure it gives you in
making it. With foresight, careful planning, and proper care you can be
assured that your collection will be studied and enjoyed long into the
future, and that you have made a real contribution to our growing knowl-
edge of Lepidoptera.

THE IMMATURE STAGES OF SCOPULA ANCELLATA (HULST)
(GEOMETRIDAE)

W. C. McGuFFIN

Forestry Branch, Canada Department of Fisheries and Forestry, Ottawa, Ontario

While working in the Hedley, British Columbia, area in 1967 I collected
several adults of Scopula ancellata (Hulst) between July 4 and 12. One of
these moths laid 112 eggs. As in other species of Scopula (McGuffin 1967 )
the eggs were laid loosely on the floor of the cage or on pieces of dead
grass scattered over the floor of the cage. After eight to 10 days the eggs
hatched. The first-instar larvae accepted the foliage of white sweet clover,
Melilotus alba Desr., Chinese elm, Ulmus pumila L., and mountain alder,
Alnus tenuifolia Nutt. Some of the larvae completed development and
pupated in about 40 days. From these pupae, six males and two females
emerged September 4 to 17, 1967. The great majority of the larvae how-
ever, ceased feeding towards the end of the summer. They were buried in
glass containers in the soil on September 30, 1967, and dug up on May 12,
1968. Most of the larvae were dead when examined but a few survived.
Irom these a male and a female emerged June 27, 1968.

DESCRIPTIONS OF IMMATURE STAGES

Egg: Longer than wide, with longitudinal ridges and cross striae. Whitish to light
brown when first laid, later with red spots. Length, 0.75-0.80 mm; width, 0.34—0.50
Mim.

Larva: First instar. Length, 3-4 mm; width, 0.2-0.4 mm. Head: Width,
0.28-0.34 mm; brown. At higher magnification (100) herring-bone pattern evi-
dent on parietal lobes. Body Jight brown, with dark-brown dorsum and venter (Fig.
1). Second instar. Length, 5-8 mm; width, 0.4-0.5 mm. Head: Width, 0.36—0.40
mm; light brown, with fine brown markings in herring-bone pattern on lobes. Body
light brown, with dark brown dorsum and venter; small brown patch on Al1—A5,
inclusive, between setae 1.2 and SV2. Third instar. Length 9-10 mm; width, 0.5

mm. Head: Width, 0.48-0.50 mm. Colour pattern of head and body as in second

VoLUME 24, NUMBER | OD

os ie
fra ie

woe peg ce Feats we
ste WE ere ene Fe

Figs. 1 and 2. Scopula ancellata (Hulst), setal and colour patterns on second
abdominal segment of larva. 1, First instar; 2, mature larva.

instar. Fourth instar. Length, 12-18 mm; width, 0.7-1.0 mm. Head: Width,
0.65-0.67 mm. Light grey with brown markings in herring-bone pattern on lobes.
Body light grey, with wide dark-grey middorsal stripe and narrow addorsal line
on thorax and posterior abdominal segments; these two lines almost fused on
anterior abdominal segments to form a solid stripe. Midventral line light grey; sub-
ventral region grey. A small brown spot, anterior to seta L3 and between setae L2
and SV2, on each anterior abdominal segment. Thoracic legs light grey; prolegs
grey, with a light-grey line along each. Plates concolorous. Fifth instar. Length,
18-22 mm: width, 0.8-1.0 mm. Head: Width, 0.84 mm. Colour pattern of head
and body much as in fourth instar. Sixth instar. Length, 19-25 mm; width, about
1.0-1.5 mm. Head: Width, 1.34-1.50 mm. Light brown with brown herring-bone
pattern on lobes. Antennae light brown. Body light brown with brown middorsal
line. Brown spot between setae L2 and SV2 on Al1—A5, inclusive. Venter brown,
with light brown midventral line (Fig. 2). Legs, prolegs, and plates concolorous.

Pupa: Brown, fusiform, much as in Scopula inductata (Guenée) (McGuffin 1967,
Fig. 84e, f).

REMARKS
The mature larva of S. ancellata can be separated from the larvae of
other species of Scopula on the basis of the brown spot on the anterior
abdominal segments between setae L2 and SV2. Apparently no other
species of Scopula has this spot.
LITERATURE CITED

McGurFin, W. C. 1967. Guide to the Geometridae of Canada (Lepidoptera). Part
1. Subfamily Sterrhinae. Mem. Ent. Soc. Can. No. 50.

56 JoURNAL OF THE LEPIDOPTERISTS’ SOCIETY

GENERIC NOTES ON TWO HAIRSTREAKS NEW
TO THE UNITED STATES (LYCAENIDAE)

Harry K. CLENCH
Carnegie Museum, Pittsburgh, Penna. 15213

In the paper that follows this one, Mr. Roy O. Kendall reports the
capture in Texas of three species of hairstreaks that are well known in
Mexico, but had not previously been found in the United States. One of
these is in good taxonomic order, but the other two require revision to
bring their nomenclature up to date.

Ocaria Clench, new genus
Type species: Thecla ocrisia Hewitson 1868

Antennae composed of about 30 segments, of which the last 13 comprise the club;
four terminal segments scaleless; an additional seven ventrally scaleless; longest shaft
segment about 3.8 times as long as average club segment, the shaft slender, each seg-
ment white-ringed proximally. Eyes with dense, moderately long hair. Frons with
erect scales and long, loose, erect bristles. Palpi with terminal segment long and
slender, smoothly scaled; next proximal segment with erect ventral scales but no
bristles.

Forewing with M. from near middle of cell-end, (M:-M:2)/(M:-Ms) = 0.47; a large
scent pad filling the end of the cell from one side to the other, and extending basad to
near the origin of Cuz; scent pad composed of densely packed ochreous scales with a
few fuscous scales intermixed; pad itself rimmed with densely packed modified scales
of ground color. Hindwing with no tornal cleft, virtually no tornal lobe, and with only
a short tail at Cue, shorter than apical width of interspace Cu:-Cu».

Male genitalia (Fig. 1). Uncus lobes low, rounded, laterally narrow and quadrate,
separated by a broad low median notch; falces broadly curved, practically without an
“elbow,” apically constricted but not hooked; vinculum with slight shoulder, but no
shoulder process; posterior dorsal vinculum with thickened margin; anterior vinculum
angularly produced midlaterally, associated with moderate coremata (scales about 1% as
long as penis); saccus subquadrate, barely longer than width at middle; valvae loosely
contiguous to tips, of normal length, broad to beyond middle, then abruptly narrow,
but not tapered; penis about 2.5 times as long as valvae (1.9 « valvae + saccus), with
tip slightly upturned, armed ventrally with a terminal triangular keel, proximally den-
tate, distally smooth; two apical multidentate cornuti, one subapical cornutus, not den-
tate but bluntly acuminate at its distal end.

Remarks. Ocaria is one of the few neotropical relatives so far dis-
covered of the holarctic Satyrium. It differs from all others in the Satyrium
series (Chlorostrymon Clench; Phaeostrymon Clench; Satyrium Scudder;
Chrysophanus Scudder ) in the projecting triangular shape of the ventral
distal keel on the penis, in the presence of an anterior process on the vin-
culum for attachment of the coremata, and in the doubled distal (dentate )
cornutus. The thickened posterior margin of the vinculum is found only in
Chlorostrymon of the other known genera; in all the rest this thickening
appears more or less internally as the vinculum strut.

VOLUME 24, NUMBER 1 a7

Fig. 1. Ocaria ocrisia Hewitson, ¢ genitalia. Top figure, genital capsule cut
through the right vinculum and “unrolled,” with uncus and falces at top; left vinculum,
saccus and valvae to left. Middle figure, penis to same scale. Bottom figure, apical end
of penis (enlarged to show cornuti and the serrate ventral keel). Specimen from near

Gomez Farias, 300 m, southern Tamaulipas, Mexico, 9.I. 1966 (leg. H. Clench and L.
D. Miller, CM-CUA Exp. ), Slide no. C-1149, CM.

58 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Ocaria ocrisia is the only member of the genus which I have examined.
One or two additional species (South American) may ultimately be found
to belong here as well.

Thereus Hubner

Thereus Hiibner [1819], Verz. bek. Schmett. (5): 79 (type species, by monotypy:

Papilio lausus Cramer [1779] ).

Genus ? (new genus): Clench 1961, in Ehrlich & Ehrlich, How to Know the Butter-
H meee Clench 1964, J. Res. Lepid. 2 (“1963”): 254 (type species, by original
designation: Thecla bourkei Kaye 1924). NEW (SUBJECTIVE) SYNONYMY.

Until recently, no specimen of the species lausus Cramer was available
to me for study. Its peculiar pattern resembled no Heterosmaitia I had
seen and it never even occurred to me that it might belong here. I have
now examined a specimen, and its genitalia indicate beyond any doubt
that it is congeneric despite its peculiar appearance. Because it is the type
species of the Hiibnerian genus Thereus, which has some 15 decades prior-
ity over Heterosmaitia, the latter name must fall.

Two species, neora Godman & Salvin and palegon Cramer, have been
found, just as surprisingly, congeneric as well, although fortunately neither
is involved in any serious problem of generic nomenclature. The former
heretofore was placed in the genus Aftlides Hiibner (with which its under-
side pattern agrees rather well). The two species bear no pattern re-
semblance to each other whatever, nor do they resemble lausus, nor are
they at all similar to any of the other members of the genus so far as now
known.

In contrast to these species (neora, palegon, lausus ), which are so highly
dissimilar, other species (thoana, guadala, brescia, bourkei) are so similar
to one another that they pose major problems of identification. These
“cryptic” species are so extremely similar, particularly in their underside
patterns, that they possibly form some sort of mimetic association.

In my earlier paper (1964) I divided the genus into two groups. These
groups (characterized in that paper), with the species now known to
belong to them, are:

1. oppia group. Includes oppia Godman & Salvin (Middle America )
and neora Godman & Salvin (Middle America). Besides being very dif-
ferent in pattern, these two differ structurally enough to warrant even-
tually being placed in two subgroups.

2. bourkei group. The subgroups I proposed (1964) must now be re-
vised to accommodate the new additions.

Subgroup A. Male scent pad simple; corematal process broad and

triangular; larger cornutus with many teeth; valvae divergent from mid-

VOLUME 24, NuMBER 1 59

dle and of normal length (reaching about to falcal shoulder). Includes
bourkei Kaye (Jamaica), guadala Schaus (Middle America), brescia
Hewitson (neotropical, widespread), palegon Cramer (neotropical,
widespread ).

Subgroup B. Male scent pad duplex, but without a rim of modified
scales; corematal process broad, parallel-sided, long and _ apically
rounded; larger cornutus with many teeth; valvae of normal length,
contiguous to near tips. Includes thoana Hewitson (Middle America )
and possibly several South American species of similar appearance
(stagira Hewitson; erenea Hewitson ).

Subgroup C. Male scent pad duplex and rimmed with enlarged, densely
packed scales of ground color; corematal process elongate, tapering-
triangular; larger cornutus without apical teeth; valvae extremely
elongate-attenuate (similar to those of the genus Allosmaitia Clench
1964), reaching about to the tip of the falx, loosely contiguous to tips.
Includes lausus Cramer (neotropical, widespread ).

THREE HAIRSTREAKS (LYCAENIDAE) NEW TO
TEXAS AND THE UNITED STATES

Roy O. KENDALL
135 Vaughan Place, San Antonio, Texas 78201

Hurricane “Beulah” struck the mainland of extreme south Texas near
Brownsville, 20 September 1967. Thousands of acres in the Lower Rio
Grande Valley were completely inundated. Santa Ana National Wildlife
Refuge located on the Rio Grande near Alamo, Hidalgo County, Texas,
was such an area. This is a favorite collecting spot for lepidopterists and
a number expressed concern over the insect life there. With so much
water, some collectors thought the insect life would be largely destroyed.
When collecting trips could be resumed, I found that little if any damage
to the insect populations had been done. On the contrary, in many ways
improvements had occurred.

Extensive flooding had germinated seeds of native plants which had
been dormant for a long time. A profusion of vegetation was produced
the following year. “Beulah” also evidently distributed insects over
wide areas, extending normal ranges for many species. This is evident
by new records in 1968 for Hesperiidae, Lycaenidae, and Heliconiinae

60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

from Texas and the United States. Only the Lycaenidae are treated in
this paper.

It will be interesting to see if these species become established in the
Refuge, and if so for what period of time. A number of species found in
Texas appear to be periodic residents when conditions are favorable.

Ocaria ocrisia Hewitson, 1865

Thecla ocrisia Hewitson 1868, Descr. New Species Lycaenidae, p. 5 (TL: Ecuador);
ibid. 1869, Ill. D. Lep. Lycaenidae, p. 123, pl. 48 figs. 235, 236; Godman & Salvin
1887, Biol. C.-Amer., Lep. Rhop. 2: 49, pl. 54 figs. 5, 6; ibid. 1901, op. cit. (suppl.),
p. 718; Draudt 1920, in Seitz, Gross-schmett. Erde 5, p. 775, pl. 152 h; Hoffmann
1941, An. Inst. Biol. (Mexico) 11: 711; Comstock & Huntington 1962, J. New York
Ent. Soc. 70: 39 (Note: original description erroneously credited to “Specimen of
a Catalogue of Lycaenidae in the British Museum’ ).

Ocaria ocrisia: Clench 1970, J. Lepid. Soc. 24: 56, 58.

One 2, very worn, Santa Ana Nat. Wildlife Refuge, nr. Alamo, Hidalgo
Co., Texas, 11—xi-1968 (leg. R.O.K.).

In Mexico this is a widespread but uncommon and rather local species
occurring mostly in mesic to moist forest. It has been found (Hoffman,
l.c., and Carnegie Museum) as far north as southern Tamaulipas and
Jalisco.

Thereus palegon Stoll, 1780

Papilio palegon Stoll 1780, Pap. Exot. 3: 159, pl. 282 figs. C, D (TL: “Sierra Leona,
sur la Cote dor de Afrique,” which is erroneous; the type was probably taken in
Surinam ); Comstock & Huntington 1962, J. New York Ent. Soc. 70: 100.

Papilio myrtillus Stoll 1784, Pap. Exot. 4: 178, pl. 380 figs. B, C (TL: “Suriname” );
Comstock & Huntington 1961, J. New York Ent. Soc. 69: 176.

[Name has sometimes been misspelled mytillus. |

Thecla juicha Reakirt 1866, Proc. Acad. Nat. Sci. Philadelphia: 338 (TL: near Vera
Cruz, Mexico); Comstock & Huntington 1961, J. New York Ent. Soc. 69: 56. [Name
has sometimes been misspelled juica. |

Thecla palegon: Hewitson [1867], Ill. D. Lep. Lycaenidae, P. 86; Godman & Salvin
1887, Biol. C.-Amer., Lep. Rhop. 2: 37; ibid. 1901, op. cit. (suppl.), p. 716; Draudt
1919, in Seitz, Gross-schmett. Erde 5: 761, pl. 150 d, e; Hoffmann 1941, An. Inst.
Biol. (Mexico) 11: 707.

Thereus palegon: Clench 1970, J. Lepid. Soc. 24: 58, 59.

One 4, condition fairly good, Santa Ana Nat. Wildlife Refuge, nr.
Alamo, Hidalgo Co., Texas, 9-xi-1968 (leg. R.O.K.).

A common and widespread species of the New World tropics, known
in Mexico (Hoffmann, I.c., and in Carnegie Museum) as far north as

southern Tamaulipas. It is found chiefly in tropical and subtropical forest,
including nearby open areas.

Allosmaitia pion Godman & Salvin, 1887

Thecla pion Godman & Salvin 1887, Biol. C.-Amer., Lep. Rhop. 2: 56, pl. 54 figs. 28—
1) (TL: Duenas (Polochic Valley) and Calderas, both in Guatemala, were listed by

VOLUME 24, NUMBER 1 61

the authors; Comstock & Huntington (1962, infra) restricted it to the former); ibid.
1901, op. cit. (suppl.), p. 718; Draudt 1920, in Seitz, Gross-schmett. Erde 5: 780, pl.
155 g: Hoffmann 1941, An. Inst. Biol. (Mexico) 11: 712; Comstock & Huntington
1962, J. New York Ent. Soc. 70: 115.

Allosmaitia pion: Clench 1964, J. Res. Lepid. (1963) 2: 255.

One 2, worn, Santa Ana Nat. Wildlife Refuge, nr. Alamo, Hidalgo Co.,
Texas, 11—-xi-1968 (leg. R.O.K.).

This is an uncommon species in Mexico, occurring chiefly in scrub and
low forest, particularly in montane areas. Hoffmann (l.c.) records it from
no farther north than Tabasco and southern Veracruz. There are speci-
mens in Carnegie Museum, however, from as far north as Sinaloa (19 mi
FE, Concordia ) and Hidalgo (7 mi N Zimapan, 1830 m).

ACKNOWLEDGMENT

To Harry K. Clench, Carnegie Museum, Pittsburgh, Penn., I wish to ex-
press my sincere appreciation for determining these specimens, reviewing
this paper, providing the references cited, and furnishing additional dis-
tribution data for each species.

INEXPENSIVE PHOTOMICROGRAPHY

JoHN M. KOLYER
55 Chimney Ridge Drive, Convent, New Jersey, U.S.A.

INTRODUCTION

In essence, photomicrography usually consists of positioning a film
several inches from the ocular lens of a compound microscope to receive
the magnified image of the subject. A special camera, without lens, gen-
erally is used for this purpose; detailed descriptions are given in texts such
as those by Allen (1941) and Shillaber (1944). An inexpensive camera
may be used without removing the lens ( Loveland, 1943), but this method
has serious disadvantages, e.g. tendency of a “flare spot” (bright area in
the center of the field) to appear in the picture.

Following is a brief description of construction and operation of home-
made equipment of the camera-without-lens style using both negative-
positive (conventional ) and Polaroid processes.

CONVENTIONAL FILM
Apparatus.—The apparatus (Figure 1) consisted of two parts: (1) a
wooden base, with generous working area, on which two rigid uprights

62 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

: ee d
Fig. 1. Conventional-film camera, with 4 x 5 in. cut film holder in place, attached
to 40-1000 binocular microscope (Jewell Optical Co. ).

were mounted, and (2) a camera body, constructed of plywood, with an
attached bellows (an inexpensive item purchased from Edmund Scientific
Co., Barrington, New Jersey ). The bellows allowed variation in magnifica-
tion by changing the lens to film distance, which fortunately can be varied
widely (actual focussing is done with the microscope adjustments ). The
camera body was mounted in a rectangle of % in. plywood with slots
permitting attachment to the angle irons of the uprights using bolts and
wingnuts (see Figure 2). The camera was painted flat black to minimize
internal reflection and was designed with a hinged back (see rear view,
Figure 2) to accommodate a 4 X 5 in. cut film holder as previously found
suited for close-up (low magnification ) photography of mounted Lepidop-
tera (Kolyer, 1965). Because the microscopes at hand all had eyepiece
housings inclined at 45°, the camera was mounted at this angle.

To link the bellows to the microscope, a sleeve of black felt attached to
the bellows was slipped over the eyepiece housing and held in place by
wrapping with a piece of heavy copper wire. This method was found

VoLUME 24, NUMBER 1 63

os

Fig. 2. Rear view of apparatus of Figure 1, showing hinged back open (film holder
removed). K

advantageous because the microscope was touched only by the felt, pre-
venting marring of the instrument, and the connection was flexible so
that microscopes with a movable tube (as opposed to the movable stage
on some models) could be focussed while viewing the image on the
ground glass.

A less-sturdy but satisfactory apparatus, so light in weight that it may
be fixed to the microscope without additional support, can be constructed
from cardboard (Anonymous, circa 1958). In fact, any design or ma-
terials are suitable provided that the film is held in the focal plane.

Procedure (black & white ).—A rectangle of frosted glass was mounted
on a 5 mm thick composition board frame so that, when inserted in place
of the film holder, the frosted surface was at exactly the same distance
from the eyepiece as the film. This was used to focus the image and then
was replaced by a cut film holder. A film was exposed, and the illumina-
tor was turned on for the desired exposure time (using the switch at lower
right of Figure 1).

Excellent results were obtained with Kodak Plus-X Pan professional
Film, Estar thick base, 4 X 5 in., developed according to the manufac-
turer's instructions. Using a Tasco 16x stereo microscope (shown, with a
different camera, in Figure 3), with the Bausch & Lomb illuminator shown
in Figure 3 set at highest intensity and positioned with the illuminator
lens 4 in. from the specimen, optimum exposure time was 20 sec. The dis-

64 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fig. 3. Polaroid Model 220 camera, adapted for photomicrography, attached to a
16% stereo microscope (Tasco). Also shown is the Bausch and Lomb micro lamp
( Nicholas illuminator ) used in the examples in the text.

tance from microscope eyepiece lens to film surface was about 7.8 in.
With the Jewell binocular microscope (Figure 1) at 1000 (10x wide-
field eyepiece, 100 objective (N.A. 1.25) immersed in oil), longer ex-
posures, e.g. 2 min., were required.

Prints (4 X 5in.) were prepared with Kodak Azo F-3 paper by the con-
tact method, using a relatively long distance (16 in.) between negative
and light bulb (250 watts) to secure even light intensity over the whole
print.

Procedure (color)—Kodak Ektachrome Color Reversal Film (for
artificial light), Type B, 4 x 5 in., was used to make transparencies. De-
velopment was done at home with a Kodak Ektachrome film processing
kit, E-3, one-gallon size. Because this film was more light-sensitive than
the Plus-X Pan, exposure times were shorter, e.g. 1 sec. with the Tasco
16% microscope and illuminator positioned as above. With a 100
microscope, using a 10 objective (0.25 N.A.), more light was required.

VOLUME 24, NUMBER 1 65

Fig. 4. Black spot on the forewing of Pieris rapae (L.) ( 2 ), photographed with the
apparatus shown in Figure 3.

A typical exposure time was | sec. with the B & L illuminator tilted 15°
down from the horizontal and the lens of the illuminator only 1.3 in. from
the subject.

PoLAROID FILM

Apparatus.—A holder for 4 X 5 in. Polaroid film sheets is available
which is dimensionally the same as a standard 4 X 5 in. cut film holder and
will fit the apparatus described above, but this lists for about twice as
much as the less-expensive Polaroid cameras. Also, the 4 < 5 in. Polaroid
film is about twice as costly per photograph as the popular 3% x 4% in.
black & white Polaroid film sold in packs of 8 (Type 107, ASA 3000).
Therefore, a Polaroid Model 220 camera, intended to use both black &
white and color film, was adapted by removing the lens and replacing
with a wooden block with sliding aluminum insert which closed the open-
ing between exposures and was drawn up to open the light path when
photographs were being taken (as shown in Figure 3). The distance from
eyepiece to film was about 8 in. Again, a frosted glass was used for focus-
sing.

Procedure (black & white).—After focussing the image, the frosted
glass was removed and replaced by a film pack (Type 107, ASA 3000).
Successive photographs were made to optimize exposure time by trial

66 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 5. A black scale from the spot shown in Figure 4, mounted in Permount
(Fisher Scientific Co.) and photographed with the camera shown in Figure 3 attached

to the microscope (at 1000) shown in Figure 1. Localization of the pigment is
suggested.

and error. Examples of satisfactory pictures are shown in Figures 4 and
5. In comparison to the above-described conventional film, the Polaroid
Type 107 film is very fast (light-sensitive ) so that less light is needed. In
the case of Figure 4, illumination was provided by a Tensor lamp (G. E. 93
bulb, at high intensity ) set at 45° and 7 in. from the subject; the Tasco
microscope was used, as shown in Figure 3. Using a stopwatch, optimum
exposure time was about 1.5 sec. (1 sec. was too short and 3 sec. too long
for best intensity in the photograph). For Figure 5, the Jewell binocular
microscope was used at 1000X as in a preceding example. An exposure
time of 2-3 sec. was suitable.

Procedure (color).—The preceding procedures were repeated using a
Polariod color film pack (ASA 75), 3% x 4% in. prints (8) as with the black
& white. Since this is a slower film than the black & white 3000 speed, longer
exposure times were necessary. With the 16x stereo microscope (B & L
illuminator at 45° with illuminator lens 4.5 in. from subject), an exposure

VoLuME 24, NUMBER | 67

Fig. 6. A white scale from a wing of P. rapae which had been exposed to chlorine
gas to stain the pterin pigment particles by murexide formation. This is a black &
white reproduction of a Polaroid color picture taken with the apparatus used for Figure

5.

time of 10 sec. was suitable. With the 1000 microscope, 1 min. was too
short (very dark print), 7.5 min. gave a somewhat dark print, and 11 min.
(used for Figure 6) was satisfactory. The colors deviated considerably
from reality but may serve to distinguish features stained differentially.
In Figure 6, here reproduced in black and white, the irregularly-shaped
particles on the scale were red-violet to the eye and appear dark violet
(against a pale blue-green background ) in the photograph.

CONCLUSION

A photomicrographic camera using conventional films may be home-
made and give excellent results, but the inexpensive Polaroid cameras now
available (color models for under $30) are readily adapted for photo-
micrography and offer the advantage of “instant” pictures using the
popular, relatively low-cost film packs. Polaroid color photographs, useful

68 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

at least in cases of differential staining, are easily taken once exposure time
has been optimized. Film cost is a little over 50¢ per color picture.

LITERATURE CITED

ALLEN, R. M. 1941. Photomicrography. D. Van Nostrand Co., Inc., New York City.
AnonyMous. Circa 1958. American Optical Co. Reports on Teaching with the
Microscope. American Optical Co., Instrument Division, Buffalo, New York.
Kotyer, J. M. 1965. An inexpensive apparatus for photographing mounted specimens.
J. Lepid. Soc. 19(4): 212-214.

LovELAND, R. P. 1943. Simplified photomicrography with a hand camera. Science
97( 2505): 24-26.

SHILLABER, C. P. 1944. Photomicrography in Theory and Practice. John Wiley and
Sons, Inc., New York City.

NOTES ON THE GENUS CEPHISE EVANS, WITH A NEW RECORD
FOR MEXICO (HESPERIIDAE)

H. A. FREEMAN
1605 Lewis Drive, Garland, Texas

When W. H. Evans (1952, p. 153) described the genus Cephise, he
made the statement that the actual systematic position of the genus was
doubtful. “The elongated wings and the conspicuous tornal lobe H indi-
cate affinity with Chrysoplectrum in the Urbanus group, while the broad
costa points to Achalarus. But the genitalia are not of the Urbanus type:
the very broad-ended uncus with a snow-white dorsal tuft is peculiar.
The proximity of veins 7 and 8 F rather than veins 6 and 7 point to the
Celaenorrhinus group near Nascus and there is a similar sexual difference,
F spots yellow in male, white in female, though not differing in size or dis-
position. The position in repose is unknown. ¢ upf with an unusually long
costal fold, reaching to beyond the discal spots. Clasp with a slender
style.”

Evans also stated that there was but a single species, cephise (Herrich-
Schaeffer) 1869, represented by two subspecies. In the nominate sub-
species the cuiller of clasp is long, narrow and straight. The spot in space
lb in both sexes is against the outer edge of the spot in space 2. The sub-
species is recorded from Honduras, Panama, Fr. Guiana, Surinam, Upper
Amazons (St. Paulo d’Olivenca), Para, Ecuador and Peru. The other
subspecies is hydarnes (Mabille) 1876, which has the cuiller of clasp broad

ind irregular. The females have the spot in space Ib on the upper surface

_ the primaries with the upper edge exactly against the lower edge of the

VOLUME 24, NUMBER 1 69

spot in space 2. This has been recorded from Matto Grosso, S. Brazil (Rio,
Sao Paulo, Blumenau) and Paraguay. Actually, cephise and hydarnes
represent separate species on the basis of their genitalia and the spot ar-
rangement of the females.

During June 1969, while collecting on the grounds of Hotel Covadonga,
six miles south of Ciudad Valles, S. L. P., Mexico, I caught two males and
a female of cephise; this represents a new skipper record for Mexico. Ob-
servations made during this collecting period makes it possible to correctly
place systematically the genus Cephise. The three specimens collected
were found in rather dense jungle. Their flight pattern and position in
repose is very similar to members of the genus Urbanus. During the
middle of the day when the jungle is hot and still cephise rests on the
under surface of banana leaves or other similar vegetation with the wings
folded. When disturbed they dart out and fly for a short distance but
before settling again will abruptly hit the underside of a leaf two or three
times before coming to rest. This characteristic is also exhibited by mem-
bers of the genus Urbanus and Astraptes. Apparently this approach will
disturb any predatory spider which might be present; the jungle is full of
such arachnids.

From information obtained by observing the habits of Cephise cephise
I now rearrange the position of Cephise and place it between Urbanus and
Astraptes. Certain morphological as well as the behavioral characteristics
indicate that this is a much better arrangement than that of Evans in which
it follows Nascus, a member of the Celaenorrhinus group the species of
which hold their wings flat in repose.

ACKNOWLEDGMENT

I would like to express my appreciation to the American Philosophical
Society for a research grant which made this study possible.

BIBLIOGRAPHY
Evans, W. H. 1952. A catalogue of the American Hesperiidae indicating the classifi-

cation and nomenclature adopted in the British Museum. Part II. London, British
Museum. 178 pp.

70 JouRNAL oF THE LEPIDOPTERISTS SOCIETY

y
GY
Z

OBITUARY
ALEXANDR SERGEEVICH DANILEVSKI

On the 27th of March, 1969, there died unexpectedly of heart failure at his home
in Leningrad, the eminent Russian entomologist and lepidopterist, Dr. Alexandr Ser-
geevich Danilevski. Dr. Danilevski was the leading experimental insect ecologist and
physiologist of the USSR, a prominent systematist of the Lepidoptera, and an authority
on the Tortricidae or leaf-rollers. He was Dean of the Faculty of Biology and Agronomy
and professor of Entomology at the University of Leningrad.

Dr. Danilevski was born on the 4th of March, 1911, near the town of Mirgorod, in
the district of Poltava in the Ukraine. His father was an agronomist; his mother, a

chool teacher, was the granddaughter of the great Russian poet, A. S. Pushkin. Even
as a schoolboy he was attracted to natural history and became a familiar figure at the
Poltava Museum and Experimental Station. After finishing high school in 1930 he re-
ned in Leningrad to enter the Institute of Applied Zoology and Phytopathology,
here he completed his studies in 1933 with excellent marks. At that time he was
perienced faunist and systematist of Lepidoptera. He took a post at the

VOLUME 24, NUMBER 1 fil

Federal Institute for Plant Protection, where he studied the problem of food specializa-
tion in insects. His first scientific paper, which dealt with the results of his study, was
published in 1935.

In 1936, Dr. Danilevski became reader at the University of Leningrad; his connec-
tion with this University lasted throughout the rest of his life and his entire scientific
career.

During World War II he volunteered for military service and took part in the heroic
defense of Leningrad. After the war he returned to Leningrad University, first as
reader in the Department of Entomology, and since 1955 as its head. In 1967 he was
appointed Dean of the Faculty of Biology and Agronomy. Concurrently during these
years, Dr. Danilevski worked at the Zoological Institute of the Academy of Sciences
of the USSR, where he headed the Division of Lepidoptera. He maintained a very
close contact with this Institute until the last day of his life.

Dr. Danilevski’s work on Applied Entomology and Phytopathology directed his
interest to acute problems of plant protection and so paved the way for his later great
ecological work. He studied the ecological regulation of population density and of
biological cycles of pests, starting with the interrelation between pests and their food
plants. When he undertook a study of distribution and acclimatization of certain
lasiocampid species in the USSR, he viewed the problem from ecological and
physiological aspects. His analysis showed that limits of an area of distribution cannot
be explained solely by the ecological requirements of active insect stages, but that the
resting stages of the life cycle must also be taken into account. This conclusion
awakened his interest in diapause and the factors regulating it. The results of his
studies on reactivation and adaptation of stages in diapause have become fundamental
for the understanding of many problems of ecology and zoogeography.

A new step in the development of insect ecology was provided by his studies on
photoperiodical adaptations. The influence of day-length on insect development had
been discovered at the beginning of the present century, but Dr. Danilevski was the
first to realize the great importance of this aspect of the regulation of diapause, and of
other cyclic seasonal phenomena. He showed the wide occurrence of the photo-
periodic reaction in insects, demonstrated the diversity of its occurrence, and deter-
mined the geographical variability of the reaction and its dependence on temperature,
food, and other factors of the environment. He was the first to study the hereditary, as
well as many other aspects of the photoperiodic reaction. As the final result of his re-
search, he defined the ecological mechanism of the regulation of the life cycle in
insects. A theoretical conclusion of his research was his monograph and doctoral
thesis: “Photoperiodism and seasonal development of insects” (1961), for which he
was awarded a first prize by Leningrad University. This book has been published in
translation in England and Japan, and is generally recognized as one of the classic
works on invertebrate ecology.

To the end of his life Dr. Danilevski continued the study of photoperiodism and
so erected a school of ecologists. In 1968 his pupils published a symposium entitled
“Photoperiodical Adaptation in Insects and Acari,” in connection with the XIIIth
International Congress of Entomology.

He was also a talented and devoted teacher and had a large number of ardent
pupils ranging from University lecturers and students to school teachers and school
children, for all of whom he organized entomology classes.

At the same time, Dr. Danilevski was very active in the field of the systematics of
Palaearctic Lepidoptera. When still quite young and under the influence of the
Russian lepidopterist, N. N. Filipjev, he made special study of the Tortricidae or leaf-
rollers and soon became a great authority on this group of Microlepidoptera. In all, he
described over seven new genera and 60 new species, chiefly of the Tortricidae; he
was also an authority on lepidopterous larvae. He edited the late A. M. Gerasimov s
monograph on the caterpillars of the USSR and made many additions from his own

72 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

knowledge. He compiled keys to insect orders based on last larval stages, and pub-
lished a fauna of dendrophilous larvae of Lepidoptera of the USSR.

In systematics, Dr. Danilevski applied not only the classical methods but used his
fundamental knowledge of ecological data. All his purely taxonomical papers have an
ecological approach, so valuable, but so often completely lacking in the work of others.
This approach is evident in the recent (1968) monumental monograph on the tribe
Laspeyresiini of the USSR (tortricid fruit-borers), which was published by him and
his pupil V. I. Kuznetsov. This is unquestionably the best treatise on the group to
have appeared within the last hundred years.

The scientific merits of Dr. Danilevski have been acknowledged both in his own
country and abroad. He was member of the Board of the Federal Entomological So-
ciety of the Scientific Advising Committee of Leningrad University, of the Zoological
Institute of the Academy of Sciences, and of the Federal Institute for Plant Protection.
He was also a member of the Editorial Board of Entomologicheskoye Obozreniye and
of the international journal Insect Physiology.

Besides his scientific qualifications, Dr. Danilevski was a most amiable and kindly
man, a lively and interesting companion, and a warmly sociable person. His so un-
expected and much too early death cannot but be a severe loss for all his friends,
students, and colleagues as well as for lepidopterology in the USSR and abroad.—A.

DriaxonorfF, Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands.

MATING FLIGHT OF BUTTERFLIES WITH MIMETIC FEMALES
AND NON-MIMETIC MALES

Although no records were kept and it is therefore impossible to cite precise dates
and localities, the mating of the following five species has been observed with some
frequency over a good many years. In all cases the female was the active partner
and the nuptial flight took place in the afternoon, but earlier for Hypolimnas
misippus L. than for the species of Papilionidae.

Papilionidae:

Papilio polytes L. Mysore, (S. India) and Ceylon, 1939
Papilio dardanus Brown. Rhodesia and East Africa, 1940-1968
Papilio echerioides Trimen. Kenya and Ethiopia, 1940-1968
Papilio jacksoni Sharpe. Kenya Highlands, 1955-1968

Nymphalidae:

Hypolimnas misippus L. East Africa, 1955-68, Aldabra Island, (Indian Ocean,
March 1968)

The species listed above show very conspicuous sexual dimorphism, and there is
n0 possibility of error. It is also logical that these species should be protected by
exposure of the (mimetic) female pattern during the very slow and vulnerable
mating flight. It is likely that most species with mimetic females and non-mimetic

es behave in a similar fashion, but the matter requires further study.—R. H.

‘CASSON, Centennial Museum, Vancouver, British Columbia, Canada.

VOLUME 24, NUMBER 1] es

MINUTES OF THE SIXTEENTH ANNUAL MEETING OF THE PACIFIC
SLOPE SECTION OF THE LEPIDOPTERISTS’ SOCIETY

SEPTEMBER 5-7, 1969, LOS ANGELES, CALIF.

FRIDAY, SEPT. 5

The formal meetings took place at the Los Angeles.County Museum of Natural His-
tory in Exposition Park. On Friday afternoon some of the early arrivals visited the
Entomology Section in the museum building. The collections were available for study,
and as is known by many, it is especially rich in Lepidoptera; it contains many older
specimens taken by pioneer collectors in the “classic” Southern California localities. Dr.
C. L. Hocur was host, and as member of the program committee, arranged the sub-
sequent dinner, luncheon and banquet.

Later, an informal social gathering for cocktails and dinner took place at a nearby
restaurant on Figueroa St. This included a few of the later arrivals. Proceedings
gained momentum with an after-dinner “beer-bust’” at an adjoining “water-hole.”

SATURDAY, SEPT. 6

Members and guests gathered at the museum building for registration and informal
discussion. Coffee and doughnuts, compliments of the Lorquin Entomological Society,
were available.

Shortly after 10 a.m. the meeting was called to order by Dr. C. L. Hocur. R. L.
LANGSTON was unanimously elected Secretary pro tem. Dr. J. W. TILDEN was elected
chairman of the Resolutions Committee.

The Address of Welcome by Dr. HERBERT FRIEDMANN, LACMNH Museum Director
was read in his absence by Dr. Hocue.

The Presidential Address by Dr. DAviy Harpwick, Canada Dept. of Agriculture was
also read in his absence. It was stated that the 1970 meeting will be held in conjunc-
tion with the National Meeting in Carson City, Nevada.

E. J. Newcomer, Yakima, Wash., informed the group that, on the basis of the mail-
ings of the NEWS, the Society has enjoyed a 33% increase in members.

Dr. Hocur announced the retirement of LLoyp M. Martin on July 31, 1969, and
that at the time of the meetings, Mr. MARTIN was visiting museums in Europe. A
“Best Wishes” card was placed at the coffee table, to be signed by the Lepidopterists.

The Saturday morning session included two formal presentations:

Papilio hybrid studies—FRED THORNE, El Cajon, Calif.

Hand pairing of Papilios—W1LL1AM HeEnpcEs, Lakeside, Calif.

A showing of slides, with Cart GooppAsTuRE as projectionist included:

Nevada collecting spots—PETER HERLAN, Carson City, Nevada

Alaskan butterfly life—KENELM Purip, Fairbanks, Alaska (narrated in his absence
by Carl Goodpasture ).

Colias of the Palearctic compared to the Nearetie—WILLIAM Hovanitz, Arcadia,
Calif.

Collecting spots in Oregon—Warners, Steen Mtns.—E. J. NEwcoMer, Yakima,
Wash.

Speyeria leto pugatensis, gynandomorph from northwestern California—STERLING
Marroon, Chico, Calif.

“The five mountain ranges around Albuquerque” plus slides of Speyeria, Ca-
tocala, etc.—RicHARD HoLianp, Albuquerque, New Mexico.

The meeting was then adjourned for a group luncheon at the restaurant on Figueroa
St. Generous portions were served, and no long speeches occurred.

At 1:35 p.m. the meeting was called to order by Dr. J. W. TrwpEN. Mrs. CHARLES

74 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

HocvE was introduced as the hostess for the Saturday afternoon LADIES PROGRAM. The
program included a visit to the Los Angeles County Museum of Art.

The Saturday afternoon session consisted of several studies:

Distributional patterns of California moths—RoONALD LEuscHNER, Gardena, Calif.

Microlepidoptera associated with oaks in California—PavuLt Op er, Albany, Calif.

Mechanisms of sex pheromone communication among moths—Harry SHOREY,
Riverside, Calif.

A brief coffee-doughnut break ensued, then back to the tight schedule.

A collecting trip in northern latitudes and “‘Rediscovery of Pieris napi colei”—
Don Err, Boulder, Colorado (read in his absence by Dr. Tilden).

An Open ForuM on NOMENCLATURE was next on the agenda, led by the following
panelists:

Pappy McHenry, Burbank, Calif.—‘‘Introduction to nomenclature.”
E. J. Newcomer, Yakima, Wash.—‘‘Pronunciation.”’

Lively discussion followed, but was cut short because of the lateness of the hour.
Before closing, Dr. J. A. PowELL, Berkeley, Calif., gave a report on the Lepidopterists’
SociETy meeting in East Lansing, Michigan, June 16-17, 1969. The meeting was then
adjourned by Dr. TiLDEN at 5:10 p.m.

The annual banquet was held at the Cockatoo Inn in Hawthorne, several miles
down the freeway and dozens of stop-signals later. Following the sumptuous dinner,
there was a showing of the EucENE O. MurMAN transparencies. These paintings shown
through two projectors (some to indicate iridescence ) were mostly of North American
butterflies. It was ably narrated by Dr. Joun Gartu, Allan Hancock Foundation, Los
Angeles, Calif.

SUNDAY, SEPT. 7

Members and guests again gathered at the LACMNH, with the meeting being called
to order shortly after 9 a.m. by Dr. Hocue.

ELtTon O. SEtTE presided over the Sunday session, at which the following papers
were presented:

Co-existence of Plebejus in Southern California—Cari_ GooppasturE, Pasadena,
Calif.

Relationships within the acmon group of Plebejus—Dr. J. W. TiLpEN, San Jose,
@alit

A short break was then taken to partake of coffee and doughnuts, again furnished by
the Lorquin Entomological Society. The program resumed with:

Butterflies of five central New Mexico mountain ranges—RicHARD HOLLAND,
Albuquerque, New Mexico.

Notes on rearing Speyeria—STERLING MatTroon, Chico, Calif.

Evolution in the genus Argynnis—Dr. Witt1AM Hovanirz, Arcadia, Calif.

Anthocaris bionomics and distribution in the Great Basin—Rev. Davin L. BAUER,
South Lake Tahoe, Calif.

The “Collecting reports” next on the agenda were omitted because of the very
crowded schedule. However, several Lepidoptera collections by various members had
oS on wey during the two days, and reprints and mimeographed papers were
made avallaple

The business meeting was then convened by ELTon O. SETTE.

The Society Library: Dr. PETER BELLINGER, Northridge, Calif. reported on the

status of the library. lt has expanded considerably with the addition of many books
and reprints. Also on file are the field season reports. Numerous scientific journals are
received in exchange for our JOURNAL. These materials are available to all members,

to borrow either in person or by mail.

Preasurer s report: NELSON Baker, Santa Barbara, Calif., reported that $95 is on
deposit in the account. An additional $39 was received from registrations at the cur-

VOLUME 24, NUMBER 1 TE

rent meeting. However, from this, more than $12 was paid for banquet entertainment.
Another $10 was used for postage and program printing. It was moved that the
treasurer secure a permanent record book. A vote of confidence was given Nelson
Baker to continue as the Pacific Slope Treasurer, with checking account rights to be
extended to Dr. Hogue.

Dr. J. W. TrLpEN, Chairman, proposed the following resolutions:

1) That a vote of thanks be extended to Dr. Friedmann and the staff of the Los
Angeles County Museum for use of the facilities of the museum.

2) Also to Charles Hogue, Lloyd Martin, and Fred Thorne for their work in planning
the program and making arrangements for the meetings.

3) Also to the Lorquin Entomological Society for providing the coffee and snacks.

4) That the Secretary pro tem. be asked to extend to the President and Society an
invitation to meet jointly with the Pacific Slope Section in 1970 at Carson City.

5) That members of the Society and Registrants, wishing to take this opportunity of
acknowledging the contributions of Lloyd Martin to the Society and its individ-
ual members, extend their greetings to Mr. Martin on the occasion of his
formal retirement, in the hope and expectation that he will in fact continue his
activity for many more years.

6) That it is the sense of this meeting that the Lepidopterists’ Society should begin
appropriate action, as a body, and in conjunction with other organizations, to
obtain protection for endangered species of insects, other animals and plants,
and to preserve the natural ecology of North America and the rest of the World.

The first five were either passed or approved by acclamation. The sixth point on
conservation entailed further discussion.

A motion was made by Fred Thorne that a committee be appointed for the Society
to enter into the field of conservation. Paul Opler stated that the conservation issue
had already been discussed at the National meeting. Further discussion revealed that
conservation is a matter to be considered by the Society as a whole. Several people
expressed opinions for and against. Subsequently the whole matter was tabled.

It was moved by Fred Thorne that the Society congratulate Peter Bellinger on the
wonderful library work. Approval was unanimous.

Selection of 1970 meeting place and date: Paul Opler and Jon Shepard were on
the committee for merging the National meeting with the Pacific Slope meeting, and
Carson City was chosen. The location was agreeable to President Hardwick, Peter
Herlan and others concerned. Charles Harbison suggested that the last weekend in
June would be appropriate. This was not acted upon, but Herlan stated that any sum-
mer date chosen would be agreeable to the eastern group. Dr. Tilden proposed that
we accept the recommendations from the established (Opler) committee. Dr. Powell
amended the motion to accept in advance that date for the meeting ultimately decided
on by the (to-be-elected) Program Chairman. Nelson Baker moved that we choose a
definite date. The original (Tilden) motion was passed after much interim discussion.
The amendment ( Powell) was withdrawn by Powell. In summary, Noel La Due moved
that the time and place be selected by the Program Committee. The motion passed,
but most present were confused by this time. Opler moved that Herlan be in charge of
local arrangements, and that David L. Bauer and Tilden be co-chairmen of the pro-
gram committee. The motion was passed.

Selection of site for 1971 meeting: Paul Opler moved that the 1971 program com-
mittee be selected by the chairman of this session (Sette). Sette selected Thorne as
Chairman, with the latter to be assisted by McHenry and Powell. The selection was
approved.

Amid a last minute flurry of discussion and general visiting, the meeting was noisily
adjourned. A few of the lingering members had a late lunch in the museum cafeteria;
these and others also re-visited the Entomology Section.

76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ATTENDANCE

Although most delegates were from California, five other states were represented.
Members, guests and speakers who attended all or part of the meetings and banquet
included:

N. W. Baker & wife, D. L. Bauer, P. F. Bellinger, R. W. Breedlove, C. J. Callaghan
(Utah), J. F. Emmel, M. Eloas, G. S. Forbes, J. Garth & wife, C. Goodpasture, C. F.
Harbison, W. A. Hedges & wife, J. B. Heppner, P. J. Herlan (Nevada), C. L. Hogue
& wife, R. Holland (New Mexico), W. Hovanitz & wife, K. C. Hughes, W. Klein, N.
La Due & wife, J. Lane & date, R. L. Langston, R. Leuschner & wife, S. Mattoon &
wife, P. McHenry, J. R. Mori, E. J. Newcomer (Washington), P. A. Opler & wife, B.
Perkins & wife (Oregon), J. A. Powell, C. Quick, A. Rubbert & wife, C. A. Sekerman,
O. E. Sette, O. A. Shields, H. Shorey, J. S. Snider, W. L. Swisher, Fred Thorne, K.
Tidwell & wife (Utah), J. W. Tilden & wife, and M. Toliver (New Mexico).

Ropert L. LAncston (Secretary pro tem.), 3 Arlington Ave., Berkeley 7, Calif.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
IN (Gi, 88). OE

Required six-month’s rotice is given on the possible use of plenary powers by the
International Commission on Zoological Nomenclature in connection with the follow-
ing names listed by case number:

1867. Suppression of Huebner’s pamphlet Der Schmetterlinge Lepidoptera Linnaei
europdisches Heer, circa 1790-1793.

1875. Suppression of Papilio saportae Huebner, 1828/32 (Insecta, Lepidoptera).

1876. Type-species for Agrotiphila Grote, 1875 (Insecta, Lepidoptera ).

Comments should be sent in duplicate, citing case number, to the Secretary, Inter-
national Commission on Zoological Nomenclature, c/o British Museum (Natural His-
tory), Cromwell Road, London S.W.7, England. Those received early enough will
be published in the Bulletin of Zoological Nomenclature.

November 1969

W. li. Cuina, Assistant Secretary to the International Commission on Zoological No-
menclature.

a

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
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Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
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Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
heading LireraturReE Cirep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: All photographs and drawings should be mounted on stiff, white
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sity of Louisville, Louisville, Kentucky 40208.

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyr F. pos Passos

Price: Society members—$4.50, others—$6.00; uncut, unbound signatures
available for interleaving and private binding, same prices; hard cover bound,
add $1.50. postpaid

ALLEN PRESS, INC. ene LAWRENCE, KANSAS
US. m&

CONTENTS

Careasson, R. H. Mating flight of butterflies with mimetic females and

non-mimetic ‘miales)) ee 72
Clench, H. K. A new subspecies of Brephidium exilis from Yucatan (Lep-
idoptera::Lyeaenidae) 000 3- 6
Clench, H. K. Generic notes on two hairstreaks new to the United States
(Lycaenidae) (i 56-59
Covell, C. V.. What's your collection worth 000 ee 51-54
Diakonoff, A. Obituary of Alexandr Sergeevich Danilevski _ 70-72
Emsley, M. G. An observation on the use of color for species-recognition
in Heliconius besckei (Nymphalidae) ee 25
Freeman, H. A. Notes on the genus Cephise Evans, with a new record for
Mexico \{ Hesperiidae) 000000 68-69
Greene, G. L. Head measurements and weights of the bean leaf roller,
Urbanus ‘proteus (Hesperiidae) 0.0 47-51
Kendall, R. O. Three hairstreaks (Lycaenidae) new to Texas and the
United’ States ioe Ve AUS Ode 59-61
Kolyer, J. M. Inexpensive photomicrography 61-68
Masters, J. H. A new Caltithomia (Ithomiidae) from Bolivia 22-24

Masters, J. H. A new Perisama (Nymphalidae: Callicorini) from Bolivia . 10-13

Masters, J. H. Bionomic notes on Haeterini and Biini in Venezuela (Sa-

fyridae). 0 a Mn ONE ARO OSS Cy GSS ert 15-18
McGuffin, W. C. The immature stages of Scopula ancellata (Hulst) (Geo-
metridae) 00 ui OE OU TU UE IN A NS Sr 54-55

Miller, L. D. Multiple capture of Caria ino melicerta (Riodinidae ) at light 13-15
Muller, J. New records for New Jersey with notes of other scarce captures

Wr SL OGT! oo I i SES A aU Rr l- 3
Munshi, G. H. and S. A. Moiz. Studies on the biology and seasonal history

of Polydorus aristolochiae (Papilionidae) 19-22
Nielsen, M. C. New Michigan butterfly records 0) 42-47
dos Passos, C. F. A revised synonymic catalogue with taxonomic notes on

some Nearctic Lycaenidae 0 Or 26-38
Povolny, D. F. An interesting new species of the New World Gnorimosche-

mini (Gelechiidae) from the Lesser Antilles 0. 6-10
Steyskal, G. C. On the nature and use of the suffix -ellus, -ella, -ellum in

SPECIES-Zroup, NAMES oO TO ta ile he 38-41
Minutes of 16th Annual Meeting of Pacific Slope Section 73-76

Zoological Nomenclature

Volume 24 1970 Number 2

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

29 May 1970

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

D. F. Harpwicx, Editor of the Journal
C. V. Covet, Editor of the News
S. A. HesseL, Manager of the Memoirs

EXECUTIVE COUNCIL

E. B. Forp (Oxford, England ), President

C. L. Remincron (New Haven, Conn. ), President-elect
L. P. BRower (Amherst, Mass.), Ist Vice President

L. M. Martin (Prescott, Ariz.), Vice President

J. W. TxxvEN (San Jose, Calif.), Vice President

S. S. Nicoay (Virginia Beach, Va.), Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

Members at large (three year term): W. C. McGurrin (Ottawa, Ont.) 1971
J. F. G. Cuarxe (Washington, D.C.) 1970 Y. NexruTenxo (Kiev, U.S.S.R.) 1971
H. K. Ciencu (Pittsburgh, Penna.) 1970 B. Maruer (Jackson, Miss.) 1972

B. Wricut (Halifax, N.S.) 1970 M. Ocata (Osaka, Japan) 1972

A. E. Brower (Augusta, Me.) 1971 E. C. Wetuinc (Merida, Mexico) 1972

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
and the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

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for February, May, August and November, and eight numbers of the News each year.

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JOURNAL OF

Tue LerpipoprtreERiIstTs’ SOCIETY

Volume 24 1970 Number 2

THE ENVIRONMENTAL REGULATION OF SEASONAL
DIMORPHISM IN PIERIS NAPI OLERACEA (PIERIDAE)

CHARLES G. OLIVER

Department of Biology, Yale University, New Haven, Connecticut

The causes of seasonal changes in the coloration, wing marking pattern,
and even wing and body structure of Lepidoptera have long been an
intriguing subject for experimentation. The changes may be of .two
basic types: either a marked but continuous shift in intensity and extent
of pattern elements or ground color, or a sharply defined dimorphism in
which colors and pattern elements may be completely replaced and
changes in wing and body structure may occur. The former type is com-
mon in continuously brooded species such as Colias eurytheme Boisduval
and Phyciodes tharos Drury, while the latter occurs most often in dis-
cretely brooded species such as Araschnia levana L. and Celastrina
argiolus L.

Two chief factors, temperature and photoperiod, have so far been
shown to be of importance in regulating seasonal forms. In Precis octavia
Cramer the effect is caused entirely by rearing temperature (McLeod,
1968). The appearance of female Ascia monuste L., which may vary in
color from white to dark gray (Pease, 1962) and the seasonal dimor-
phism of Pieris protodice Boisduval & LeConte (Shapiro, 1968) are, on the
other hand, due to daily photoperiod exposure during the immature
stages. The forms of other species such as Araschnia levana (Siffert,
1924; Miller, 1955, 1956) and Colias eurytheme (Ae, 1957) apparently
are regulated by a synergism between temperature and photoperiod.

Pieris napi oleracea Harris is sharply bivoltine in eastern North
America. The spring and summer broods differ markedly in body to
wing size proportions and in the extent and intensity of wing pattern
(Table 1). The experiments described here were designed as an attempt
to produce the seasonal forms shown in nature using temperature and
photoperiod as controlled variables.

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Differences in appearance between “spring” and “summer” phenotypes
of Pieris napi oleracea.

“Spring” phenotype “Summer” phenotype
1. Body relatively stout, dark, hairy. Body relatively slender, lighter-colored,
smooth.

2. Dorsal dark scaling at wing bases and Dorsal dark scaling light.
forewing apex heavy.

3. Scaling along veins on underside of Scaling along veins on underside of hind
hind wing always very dark, complete, wing varying from light and complete, to
clear-cut. light and incomplete, to absent.

Procedure

Four broods of P. n. oleracea were derived from wild-inseminated
females collected 15 May 1968 at Wahconah Falls State Park, Berkshire
Co., Massachusetts. The newly hatched larvae of three of the females
were divided into two lots for rearing both on long days, short nights
(15hL/9hD ) and on short days, long nights (12hL/12hD), while those
of the fourth female were reared only on long days, short nights. All of
the larvae were fed on cut mustard leaves (Brassica) and given a 25° C
day and a 22° C night.

None of the pupae produced from long-day larvae entered diapause,
and are hereafter referred to as the non-diapausing pupae. Among these
emergence or death occurred in every case within six to ten days. None
of the pupae produced from short-day larvae had emerged after two to
three weeks and were thus presumed to have entered diapause (the
diapausing pupae). They were placed at 5° C in darkness for 10 weeks
to terminate diapause.

Immediately after pupation the non-diapausing pupae were subdivided
into four lots and redistributed to await adult development under the
following temperature and photoperiod regimes: 25° C day, 22° C night,
I5hL/9hD (N=14); 25° C day, 22° C night, 12hL/12hD (N =13);:
25° C day, 15° C night, 16hL/ShD (N=12);, 25° € day, 5 °a@enmente
16hL/8hD (N=18). Mortality of the pupae was 8/65 (12.3%), due
probably in some cases to cold shock.

The diapausing pupae were left on their larval temperature and photo-
period regimes until the onset of chilling. After removal from cold, they
were subdivided into three lots, all kept on long days and at 25° C during
the day, but with night temperatures of 22° C (N = 14), 15° C (N =8),
5° C (N=8). Hatching of 30/43 (69.8%) of the pupae occurred six
to 18 days after removal from cold. The death of another pupa occurred
before 18 days after chilling, while the other 12 pupae were kept at

VoLUME 24, NuMBER 2 79

‘A,

Figs. A-D, Pieris napi oleracea, ventral view of males. A, “Spring” adult from
diapausing pupa; B—D, extreme light, medium, and extreme dark “summer” adults
from non-diapausing pupae. (All laboratory-reared progeny of wild-caught females
from Berkshire Co., Mass., 15 May 1968. )

room temperature for several months and gradually desiccated without
ever hatching.

Results

The results were unequivocal. In each case the non-diapausing pupae
produced adults typical of the “summer” phenotype, while the diapausing
pupae produced typical “spring” adults. There was no correlation of
phenotype to any environmental factor except larval photoperiod ex-
posure. Each of the four broods gave the same range of adult variation
in coloration as found in the wild summer emergence (Table 1 and Figs.
C and D). The diapausing pupae gave adults all almost identical in

80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

appearance to each other, regardless of pupal photoperiod and tempera-
ture regime (Figs. A and B).

Discussion

Clearly the basic environmental factor determining seasonal form in
P. napi oleracea is larval photoperiod exposure. However, it is impossible
as yet to say whether it is larval photoperiod itself or the occurrence
of diapause in the pupa that actually directly determines which phenotype
is to be produced, since larval photoperiod and diapause cannot here be
separated.

It is interesting that even though part of the non-diapausing pupae
were chilled overnight during the time when adult wing pigment was
being formed, there was no darkening effect at all shown in the emerging
adults. This seems to be in contradiction to Merrifield’s (1893) report
that chilling summer pupae of British P. napi L. caused the emerging
adults to have “most, but not all, of the characteristics of the spring form.”
Merrifield’s pupae were, however, chilled for three to four months prior
to exposure to spring (average 54° F') or summer (average 80° F) con-
ditions during development. There was no further intensification of dark
markings in the lot exposed to spring conditions. Probably the pupae had
entered diapause at the onset of the three or four months of chilling, but
it may be that P. n. napi has a rather different system for the regulation
of seasonal forms from that of P. n. oleracea.

The results indicate that P. n. oleracea in eastern North America has
two systems that regulate the appearance of the adult phenotype. The
first is the inherited, environment-independent expression of dark wing
markings in the “summer” form. This system is responsible for the indi-
vidual variation found in the summer brood. The second is the environ-
mentally induced “switching on” of the “spring” form. Since this involves
uniformly maximum expression of the characters that show variation in
the summer form, as well as new differences, any genetic variation in
wing pattern that would be revealed in the summer brood is concealed
in the spring brood.

Summary

Diapausing and non-diapausing pupae of Pieris napi oleracea were
exposed to various photoperiod and temperature regimes during adult
development. The pupae which had undergone diapause produced only
adults of the “spring” form, whereas those that had not, produced only
adults of the “summer” form. Regulation of seasonal dimorphism is thus

VoLUME 24, NUMBER 2 81

controlled by larval photoperiod exposure, which also controls the induc-
tion of pupal diapause.

Acknowledgment

I thank Dr. Charles L. Remington for his helpful advice during the
preparation of this paper.

Literature Cited

AE, S. A. 1957. Effects of photoperiod on Colias eurytheme. Lepid. News 11:
207-214.

McLeop, L. 1968. Controlled environment experiments with Precis octavia Cramer
(Nymphalidae). J. Res. Lepid. 7: 1-18.

MERRIFIELD, F. 1893. The effects of temperature in the pupal stage on the colouring
of Pieris napi, Vanessa atalanta, Chrysophanus phlaeas, and Ephyra punctaria.
Trans. Ent. Soc. Lond. 41: 55-67.

Mtuier, H. J. 1955. Die Saisonformbildung von Araschnia levana, ein photo-
periodisch gesteurter Diapause-Effekt. Naturwiss. 43: 134-135.

1956. Die Wirkung verschiedener diurnaler Licht-Dunkel-Relationen auf
die Saisonformbildung von Araschnia levana. Naturwiss. 43: 503-504.

PEASE, R. W. 1962. Factors causing seasonal forms in Ascia monuste (Lepidoptera).
Science 137: 987-988.

SHAPIRO, A. M. 1968. Photoperiodic induction of vernal phenotype in Pieris pro-
todice Boisduval & LeConte (Lepidoptera: Pieridae). Wasmann J. Biol. 26:
137-149.

SUFFERT, F. 1924. Bestimmungfactoren des Zeichnungsmuster beim Saison-Dimor-
phismus von Araschnia levana-prorsa. Biol. Zbl. 44: 173-188.

A PLASTIC RELAXING BOX FOR PINNED AND
PAPERED SPECIMENS

ROBERT S. ROZMAN
9704 Belvedere Place, Silver Spring, Maryland

This relaxing chamber can be readily made by anyone with average
mechanical ability for just a few dollars. The setup offers several ad-
vantages over regular relaxing chambers. First, pinned specimens can be
readily relaxed without the rapid rusting resulting from pins being stuck
directly into wet sand. Second, envelopes are not in direct contact with
the wet sand. Although specimens take slightly longer to relax, they do
not become soaked and ruined. Third, the clear plastic enables one to
observe the conditions of the specimens during relaxation without open-
ing the box.

The box and lining strip are of clear plastic, such as Plexiglas, available
from hobby shops in “46 inch thickness. Sections can be bonded together

82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

LID

3/16"

SCREEN

RENO:
IDEANE

BOTTOM
SCREEN

OSS OSINCS NOD OD

CURVED
EDGE

’
33/16" :
aot Nees
<n
Fig. 1. Details of relaxing box and rack.

with a cement such as Duco Cement which will fill up any spaces formed
by irregular sawing techniques. This will form watertight joints. If all
edges are made planar and properly perpendicular, a neater bond can
be made by injecting acetone or other plastic solvents along the seams.

VoLUME 24, NuMBER 2 83

Details of the box are given in Figure 1. Of course, the dimensions may
be changed as desired, as long as the resultant base is watertight. As can
be seen, the beveled inner strip serves to hold the lid securely as in a
regular insect box or drawer.

About % inch of wet sand is placed on the bottom and a teaspoonful
or so of chlorocresol sprinkled over the sand to prevent mold (Tindale,
1961). A rack to hold the specimens is laid over this (Fig. 1). The frame,
held together with Duco Cement, is made of polyurethane foam of the
sort sold as Christmas decorations. The two screens are regular aluminum
screening available at any hardware store. These are cemented to the
top and center of the frame. In use pins are held upright by the two
layers of screening or envelopes are laid over the frame.

I have been using this type of chamber and rack for some time now.
Even fragile specimens don’t usually break. The original screening is still
shiny, the plastic still clear and the seams still watertight. I have yet to
get mold or infestation, even without prior fumigation.

Literature Cited

TINDALE, N. B. 1961. The chlorocresol method for field collecting. J. Lepid. Soc.
15: 195-197.

A REDESCRIPTION OF STRYMON BOREALIS LAFONTAINE!
(LYCAENIDAE)

J. D. LAFONTAINE
916 Innswood Dr., Ottawa, Ont.

Satyrium boreale (Lafontaine) new combination
Strymon borealis Lafontaine, 1969. Trail and Landscape 3: 151.

Upper surface of both sexes dark blackish-brown with no trace of an orange spot
at the anal angle of the secondaries. Male stigma elliptical and light grey.

Undersurface of the male slightly lighter than upper surface. Postmedial band of
primaries usually broken into three bands, separated by veins Ms and Cuz. Marginal
side of the postmedial band lined with white; basal side with very little or no trace
of white edging. Subterminal line of primaries usually reduced to only two spots
between veins Mz and Cus. The anterior three and the posterior spots of the sub-
terminal line present in falacer (Fig. 5) and caryaevorum (Fig. 6) usually completely
absent; occasionally very faint traces of these spots visible; if so, the posterior spot
with a trace of white only, never with any trace of black. Secondaries of the male

1 The name was inadvertently validated in Trail and Landscape. A _ redescription therefore
seems appropriate.

84 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Figs. 1-4. Satyrium boreale Lafontaine. 1, Underside of male lectotype. 2, Upper
side of male lectotype. 3, Underside of female. 4, Upper side of female.

Fig. 5. Satyrium falacer (Godart), underside of male.

Fig. 6. Satyrium caryaevorum McDunnough, underside of male.

with postmedial band irregular as in falacer. Subterminal line usually reduced or
absent above vein Mz.

Undersurface of female (Fig. 3) similar to male (Fig. 1). Postmedial band of
primaries more usually separated into distinct spots than in male. Subterminal line
not so reduced as in male but clearly reduced, especially so on posterior (sixth) spot
of this line. Secondaries similar to those of male but subterminal line not so reduced.

Wingspread 25 to 29 mm.

Male genitalia (Fig. 7) like those of falacer except for shape of saccus. Saccus of
boreale narrowing evenly throughout its length; that of falacer strongly constricted
subbasally giving it the appearance of an inverted bell (Fig. 8).

Lectotype here designated: 4 Britannia Park, Ottawa, Ont., July 4, 1966, J. D.
Latontaine. No. 11077 in the Canadian National Collection, Ottawa.

VoLUME 24, NUMBER 2 85

Fig. 7. Satyrium boreale Lafontaine, male genitalia.
Fig. 8. Satyrium falacer (Godart), saccus of male genitalia.

This species can be separated from caryaevorum by the incomplete
subterminal line and the much more obscure postmedial band. It is very
similar to falacer but can be separated by the subterminal line of the
primaries. A specimen of boreale which has a trace of all six subterminal
spots can be separated from a specimen of falacer with an unusually re-
duced subterminal line, by the posterior spot of this line. In falacer this
spot has a trace of black but in boreale there is only a speck of white with

86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

no black present. The subterminal line of the females of both species is
better defined than in the males. The subterminal line in a boreale female
is often almost as well defined as in a falacer male.

Boreale is usually collected on or around ash (Fraxinus), and Acer
negundo L.

Distribution: S. Que. west to Mich., south through Ohio and Penn.

Acknowledgment

I wish to thank Dr. T. N. Freeman of the Entomology Research In-
stitute, Canada Department of Agriculture, for his guidance in the prep-
aration of this paper.

A NEW SPECIES OF CAMERARIA ON BUR OAK IN MANITOBA
(GRACILLARIDAE)

T. N. FREEMAN

Entomology Research Institute, Canada Department of Agriculture, Ottawa

During the past few years some officers of the Forest Insect Survey of
Canada, Department of Fisheries and Forestry, have been studying the
biology of an undescribed species of Cameraria Chapman which is re-
ported to be abundant on bur oak, Quercus macrocarpa Michx. in the
vicinity of Winnipeg. The description of it is presented here to assist
those officers in the publication of the results of their investigations.

Cameraria macrocarpae Freeman, new species

General. Antenna white, banded with dark fuscous dorsally. Face white. Tuft
white with a few ochreous scales. Thorax golden dorsally and with a few white scales.
Forewing shining golden ochreous with three, equally spaced, outwardly angulated,
white, transverse fasciae; first at basal one-quarter, second near middle, and both
margined outwardly below angle with black scales; the third at outer four-fifths
margined with black scales throughout its length; a short, white, subapical, costal
streak followed by an area of scattered black scales in middle of wing; fringe pale
grey with a black basal line in tornal region. Hind wing light grey; fringe whitish.
Abdomen pale ochreous. Legs whitish with small black patches outwardly. Wing-
spread: 8.0-8.5 mm.

Male genitalia (Fig. 1). Uncus sub-triangular. Clasper clavate. Aedeagus clavate.
Ventral flap sub-conical.

Type material, Holotype male, Bird’s Hill, Manitoba, 10 April 1968 (reared in
laboratory). No. 11033 in the Canadian National Collection. Paratypes: one male,
Bird's Hill, 13 March 1968; four males, one female, Bird’s Hill, 22 April 1969; one

ile, Bird’s Hill, 27 April 1969 (all reared in laboratory ).

VoLUME 24, NuMBER 2 87

Fig. 1. Male genitalia of Cameraria macrocarpae Freeman.

Mine. A large upperside blotch containing several larvae.

Flight period. Late July to late August.

Remarks. This species is allied to the Myrica feeding Cameraria pic-
turatella (Braun) n. comb. It may be separated by the presence of
three, transverse fasciae on the forewing as opposed to one fascia on
that of picturatella.

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CONSIDERATIONS SUR LE MALE D’ORNITHOPTERA ALLOTTEI
ROTHSCHILD ET SUR LA PHYLOGENIE DES ORNITHOPTERES

F. Scamp
Institut de la Recherche entomologique, K. W. Neatby Bldg., C.E.F., Ottawa, Canada

ABSTRACT

Ornithoptera allottei is a species that shows a large number of characters that are
intermediate between priamus and victoriae. A detailed examination has showed that
these species do not form a progressive sequence of unilinear specialisation as it might
seem, but that priamus and victoriae have evolved in opposite directions, with allottei
representing an intermediate and more primitive condition. An explanation is given
of the gradual changes in wing patterns throughout Troides, Trogonoptera, Schoen-
bergia and Ornithoptera. A new phyletic tree of the latter is presented.

Le male d’Ornithoptera allottei a été décrit de Bougainville (Archipel
des Salomons) par Rothschild en 1914 (p. 275). En 1916, Niepelt en a
figuré les deux sexes (a: 20, pl. 17, fig. 1-2), alors que Peebless et Sch-
massmann, en 1917 (p. 426-427) en ont décrit la femelle en détail. Avec
une courte note de Niepelt parue en 1916 (b: 31) et les considérations
phyletiques de Zeuner (1943), cest-la toute la littérature qui a ete con-
Saciee a cette espece.

A tort ou a raison, Ornithoptera (Orn.) allottei passe pour étre Pespece
la plus rare du sous-genre et nous verrons dans les pages qui suivent quil
présente un interet phylétique exceptionnel. J’ai eu recemment la bonne
fortune d’en obtenir un ¢ provenant de Monoitu ( Bougainville ) et éclos en
janvier 1969. Ce spécimen mesure 149 mm. d’envergure. Sur le dessus
des ailes, la couleur métallique est bleu vert, assez terne, dans
la moitié basale des ailes antérieures et la base et le centre des pos-
térieures, alors que la moitié apicale de la bande radiale des antérieures et
tout le pourtour externe des postérieures sont teintés d'un vert jaunatre
assez vif. Le dessous est uniformément vert franc. Malheureusement, a
léclosion, Taile antérieure gauche a été fortement endommagée et une
abondante hémorragie a largement souillé les deux ailes antérieures. La
figure 1 a donc été assez fortement retouchée. Je donne aussi un dessein
des génitalia (fig. 8). Ceux de mon spécimen sont un peu différents de
la figure 54 de Zeuner: les valves sont moins longues, la téte de la harpe
non recourbée vers le bas et sa garniture d’épines hémicirculaire seulement.

Orn. allottei présente la particularité remarquable d’étre “presque ex-
actement intermédiaire” entre priamus Linné et victoriae Gray. Le Pére

! On pourait y ajouter quelques considérations parues dans “Yadoriga”’ (47, 1966: 19-20; 57,

1969: 4—6) sur les prix fabuleux atteints par cette espéce aux ventes de l’Hotel Drouot, des col-
lections Le Moult et Rousseau-Decelle.

VoLUME 24, NuMBER 2 89

‘ ,) SS

bs

Fig. 1. Ornithoptera allottei, male, légérement réduit.

Allotte qui l'a capturé le premier affirmait d ailleurs 4 Rothschild quil avait
découvert un hybride entre ces deux especes. Mais lauteur anglais publia
une note (in Peebless et Schmassmann 1917: 427) réfutant cette opinion,
a cause de l’existence de deux males identiques.

Pour commencer, nous allons examiner en détail les caractéres extérieurs
dallottei en les comparant a ceux des sous-espéces les plus primitives de
priamus et de victoriae, respectivement priamus Linné et regis Rothschild.
Nous verrons que le nombre des caractéres intermédiaires est étonnam-
ment élevé et pourtant la liste qui suit est bien loin d’étre complete. I
ny a que quatre caractéres communs avec priamus, 4, 23, 33 et peut-étre
25. Avec regis il y en a neuf, 1, 2, 3, 5, 8, 9, 16, 22 et 26, alors que tous les
autres, soient non moins de 28 sont intermédiaires.

La premiere impression qui se dégage dune telle comparaison est que
nous avons affaire a une série de trois especes présentant des caracteres
graduellement accusés et spécialisés en une “orthogénese” approximative.
De tels cas sont d’ailleurs fréquents chez les Insectes et méme la plupart
d’entre eux compremnent en général un nombre plus grand dunites. J’en
connais une belle série de huit espéces chez les Trichopteres. Mais dans
le cas qui nous occupe ici, c’est le nombre des caractéres intermédiaires qui
est etonnamment élevé.

Toutefois, il se pourrait que cette premiere impression, une série recti-
ligne de trois espéces graduellement spécialisées, soit fallacieuse et que la

90

TABLE 1.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Caractéres comparés des Ornithoptera priamus, allottei et regis.

Caracteres du corps

1) Bordure post-ocu-
laire blanche

Tache métallique
du mésonotum

Pilosité rouge des
pleures

Coloration de
Yabdomen

Bandes latérales
noires des premi-
ers segments

Faces latérales
des segments
Ill a V

6)

Face dorsale des
tergites III a VII

i)

8) Pleurites abdomi-

nales
9) Bord apical du
Ville tergite et
bord supérieur
des valves

Forme des ailes

10) Ailes antérieures

Ailes postérieures

Bord apical des
ailes postérieures

Echancrure de la
cellule I

14) Pilosité de
Vaire anale

priamus

présente

présente

presente

jaune vif

minces sur le
premier segment

uniformément jaunes

uniformément jaune

jaunes sauf les
stigmates noirs

étroitement noirs

priamus

en larges triangles
rectangles

subcirculaires, a peine
plus longues que larges
fortement crénelé

a peine indiquée

longue, brun clair

Coloration du dessus des ailes

15) Bande radiale des
antérieures

16) Bande cubitale

17) Bande anale

18) Coloration des

ailes antérieures

Coloration des
ailes

19 )

posterieures

priamus

régulierement étroite
et continue

occasionnellement
presente

régulierement étroite
et tres longue

verte, unicolore

verte, unicolore

allottei

absente
absente
absente
jaune vif

larges sur les deux
premiers segments

avec de petites
taches noires

avec une zone
décolorée

largement tachetées
de noir

largement noirs

allottei

en triangles elliptiques,
pas tres larges

1,5 fois plus longues
que larges

faiblement crénelé

assez faible

longue, beige

allottei

évanescente sur sa moitié
basale, un peu élargie
sur sa moitié apicale

absente

assez large a sa base ou
elle est unie a la bande
radiale par un sablé
métallique, puis inter-
rompue et réduite a

deux nuages intermédians

légerement bicolore,
moitié basale vert-bleu,
moitié apicale vert-jaune

faiblement bicolore,
vert-bleu au centre,
vert-jaune au pourtour

regis

absente

absente

absente

blanc grisatre

larges sur les deux
premiers segments

avec d’assez grandes
taches noires

avec une zone rosée
largement tachetées

de noir

largement noirs

regis

assez étroitement
et régulierement
elliptiques

2 fois plus longues
que larges

, LZ
non crénelé

tres nette

courte, blanchatre

regis

présente a l’extrémité
de Vaile seulement, ot
elle est tres large

absente
assez courte mais

presque aussi large
que Vaile

nettement bicolore,
verte et vert-or

fortement bicolore,
verte au centre, avec
une bande apicale
dorée

VoLUME 24, NuMBER 2

20)

21)

22.)

91

TABLE 1. Suwite.
Coloration du dessus des ailes
priamus allottei regis
Base des ailes faiblement saupoudrée nettement sablée non noircie
postérieures de noir de noir
Taches discales grandes, circulaires réduites et peu nettes exceptionnellement
noires occasionnellement présentes, nettes ou
absentes non
Cellule I des metallique noire noire
postérieures
Coloration du dessous des ailes
priamus allottet regis

23)

28 )
29)

30)

Taches discales
noires des
anterieures

Taches discales
noires des
posterieures

Taches submar-
ginales dorées

Bord noir des

postérieures

Taches marginales
noires de l’apex
des M et Cu
Tornus

Cellule et nervures
qui en radient

Coloration générale

Nervulation

31)

32)

33)

34)

35 )

Cellule des
antérieures

M3 et Cu des
antérieures

Naissance de R3

Croissant
androconial

Cellules des
postérieures

Génitalia

36)
37)

38 )
39)
40)

41)

Valves

Dents basales
de la harpe

Forme de la harpe
Col de la harpe
Téte de la harpe

Position de J’ar-
mature d’épines

en ligne réguliére dans
les cellules II 4 VIII

six, grandes et nettes

assez souvent présentes
fortement réduit

a peine indiquées

fortement jaune et noir

nettement bordées
de noir

non meétallisée

priamus

ovale, plus de trois fois
moins large que longue

longues et normalement
espacées

de la périférie de
la cellule

large, proche de
la cellule

3 fois plus longue
que large

priamus
subcirculaires

deux, peu marquées

assez large
pas indiqué

triangulaire, dirigée
vers le haut

apicale

en ligne réguliere dans
les cellules II a VIII

six plus petites et nettes

non signalées

disparu

en assez grands
triangles
faiblement jaune

cellule étroitement
bordée de noir

faiblement métallisée

allottet
aplatie 4 apex, 2,5 fois
moins large que longue

plus courtes et un peu
rapprochées

de la périférie de
la cellule

plus étroit, a mi-chemin
entre la cellule et
le bord

3,5 fois plus longue
que large

allottei
légérement ovales

une, assez gréle

étroite
peu marque

circulaire, dirigée vers
env
Varriere ou le bas

apicale et supérieure

en ligne irréguliere
dans les cellules V

a VIII

1 a 4, exceptionnelle-
ment présentes nettes
ou non

presque toujours
présentes

disparu

en grands rectangles

tres faiblement jauni

cellule a peine
bordée de noir

fortement métallisée

regis

presque cordiforme,
deux fois moins large
que longue

trés courtes et
rapprochées

de la base de R4 & 5

étroit, proche du bord
de Vaile

4 fois plus longue
que large

regis
nettement ovales

une, effilée

étirée
bien net

triangulaire, dirigée
vers le bas

circulaire

92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

situation soit plus complexe. En effet, allottei montre aussi des caracteres
communs avec croesus Wallace, alexandrae Rothschild et les Schoenbergia,
caractéres que l’on ne trouve pas chez priamus et regis. Avec croesus, al-
lottei montre des ailes antérieures en triangles assez elliptiques, a bord ex-
terne oblique et une bande anale trés courte. Avec alexandrae, allottei
présente une bande radiale étroite et nuageuse sur sa moitié basale et
dune couleur jaune-vert sur sa moitié apicale. La bande cubitale tres
courte et la bande radiale tres étroite a sa base se retrouvent aussi chez
Yensemble des Schoenbergia. Ceci suggere quallottei pourrait étre plus
primitif que priamus et que victoriae.

Pour déterminer la position et le statut dallottei, nous allons tenter de
reconstituer lévolution et la différenciation des especes constituant le
sous-genre Ornithoptera. Pour ce faire, nous passerons d’abord succincte-
ment en revue les sous-genres Troides, Trogonoptera et Schoenbergia, qui
sont plus primitifs qu Ornithoptera. Cela nous permettra de nous faire
une idée de ce quwétait ’ancétre de ce sous-genre, qui a son tour nous
servira de point de depart pour reconstituer la phylogénie de la lignée a
laquelle il a donné naissance.

Une telle étude nest pas la premiere qui ait été entreprise. En 1943,
Zeuner (p. 147) a déja analysé la phylogénie de ces lignées. Mais il s’est
place dans une perspective synchronique, cest-a-dire statique, position
aujourd/hui dépassée. L’auteur anglais a cité les caractéres communs et
les differences des diverses espéces, a évalué quantitativement leur im-
portance relative et en a déduit lordre de différenciation des espéces, qu‘il
& exprimeé en divisions arbitrairement quoique volontairement géométri-
ques (Fig. 10). Nous allons essayer ici d’aller plus loin, en nous placant
dans une perspective que la linguistique appelle diachronique. Nous ne
verrons alors plus les caractéres comme des structures seulement, mais
comme des signes et les résultats d’événements que nous tenterons de
classer en séquences et en ordre successifs, d’ou apparaitront plus claire-
ment les parentés relatives des espéces. Nous verrons alors que, quoique
les raisonnements de Zeuner soient corrects et convaincants, il y a lieu
d'effectuer des changements a son arbre phylétique. Ne connaissant pas
encore la femelle de plusieurs espéces, jai été réduit 4 utiliser les carac-
teres du male seulement, ce qui diminue malheureusement Ja valeur des
speculations qui suivent.

=

Bi ee ee

| Nervulation des ailes antérieures de Orn. priamus, allottei et regis res-
pectivement,

94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

lig. 5, 6, 7. Nervulation des ailes postérieures de Orn. priamus, allottei et regis res-
pectivement.

Troides apparait comme la lignée la plus primitive par beaucoup de
caracteres qui ont été indiqués par Zeuner. Nous n’en retiendrons que peu.
Les ailes ont une coupe primitive. Les antérieures sont noires et zébrées
de longues lignes claires, nuageuses, situées dans les cellules et longeant les

Vouume: 24, NuMBER 2 95

nervures. Ces lignes soulignent simplement les nervures et leur ensemble
ne forme pas de motifs. Les ailes postérieures sont largement jaunes
translucides et montrent primitivement six taches discales noires. Les
valve sont assez longuement ovales et la harpe bien allongée et avec une
téte nettement indiquée.

Trogonoptera vient ensuite. Aux ailes antérieures, certaines des lignes
claires du groupe précédent sont devenues vertes métalliques et sont
groupées deux a deux pour former une série de dents triangulaires ré-
gulieres, constituant un élégant motif submarginal se prolongeant en travers
des ailes postérieures qui ont enticrement perdu les écailles jaunes trans-
lucides. Les antérieures ont acquis une forme tres allongée, spécialisation
propre a cette lignée. Les génitalia ne nous seront guere utiles, car ils
sont spécialisés dans une direction tres particuliere: les valves sont plus
hautes que longues et la harpe en rectangle concave dirigé vers le bas.

Parmis les Schoenbergia, goliath Oberthiir apparait comme Tlespece la
plus archaique par lextension maximale de la zone jaune des ailes posté-
rieures. La forme des ailes est primitive. La coloration des antérieures s est
enrichie et consiste en une bande radiale, apparue soudainement, continue,
mais tres mince a sa base et une large zone cubitale et anale en triangle
rectangle. La premiere bande dérive certainement de la série de lignes
claires que certains Troides et Trogonoptera possédent contre le bord
costal, au niveau de lextrémité de la cellule. C’est pourquoi elle est
primitivement plus large a cet endroit que plus antérieurement. Son bord
interne profondément incisé de noir dans les cellules montre clairement
que la zone cubito-anale dérive, par élargissement et coalescence, dune
série de triangles submarginaux homologues de ceux des Trogonoptera. La
cellule I des ailes postérieures est noire.

Les autres Schoenbergia montrent une spécialisation en commun: la
large zone cubito-anale est maintenant plus ou moins completement
scindée en deux, ce qui fait que les ailes antérieures sont ornées de trois
bandes longitudinales: une radiale, compléte mais toujours fine ou évanes-
cente a sa base, une cubitale en large triangle aux deux extrémités effilées
et une anale courte et étroite. Les ailes postérieures ont l’aire jaune un peu
réduite mais encore importante et la cellule I reste constamment noire.
Les valves sont assez longuement ovales, caractere primitif et forment
en général deux pointes apicales inférieures, structure spécialisée.
La harpe est longue et mince, particularité primitive, mais étalée a sa base
et tres gréle a son extrémité, deux indices de spécialisation. A sa base, elle
est accompagnée de deux dents, structure primitive, qui sont devenues
gréles et barbelées par spécialisation.

Zeuner indique que la seule différence constante entre les Schoenbergia

96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 8. Valve et harpe de Orn. allottei.

et les Ornithoptera est la présence, chez ces derniers, d'un croissant an-
droconial aux ailes antérieures. Je puis donc en ajouter un second, dail-
leurs lié au premier: Schoenbergia possede une bande cubitale triangu-
laire, beaucoup plus large que les deux autres et bien distincte, sauf chez
goliath chez qui elle est encore fusionnée a la bande anale. Chez Orni-
thoptera, en conséquence de lacquisition du croissant androconial, la
bande cubitale est primitivement linéaire ou secondairement perdue. Des
ailes antérieures trifasciées sont donc un caractére primitif pour Schoen-
bergia et Ornithoptera. Il est remarquable que nous puissions suivre les
changements progressifs et continus de la coloration des ailes antérieures
depuis Troides jusque, nous le verrons 4 nouveau plus bas, chez priamus
et victoriae probleme que n’a pas pu résoudre Zeuner (p. 134).

Ceci étant posé, nous avons suffisamment d’éléments pour reconstituer
les caracteres de lancétre théorique d’Ornithoptera et en déduire la dif-
férenciation des espeéces selon le tableau des pages qui précédent.

L’Ornithoptera ancestral théorique (1, fig. 9) posséde une bande post-
oculaire blanche et de la pilosité rouge sur les pleures thoraciques. II est
probablement dépourvu de bande métallique sur le mésonotum, car cette
derniére nest présente que chez croesus et priamus, Ornithoptera moyen-
nement spécialisés et chez paradisea et meridionalis qui sont les deux
Schoenbergia les plus évolués. Ceci indique une spécialisation par conver-
gence. L'abdomen est jaune vif, assez largement teinté de noir latérale-
ment a sa base et sur les pleurites et probablement dépourvu de tache
dorsale sur les segments III et IV. Le bord apical du VIIle tergite et le
bord supérieur des valves sont largement bordés de noir.

VoLUME 24, NuMBER 2 97

Les ailes antérieures sont triangulaires et anguleuses et les postérieures
subcirculaires, avec leur bord apical crénelé et nettement échancré dans
la cellule I. La frange anale est longue.

La coloration du dessus des ailes antérieures est du méme type trifascié
que celle des Schoenbergia. Mais, 4 cause du croissant androconial qui
s étend en travers des cellules 2, 3 et 4, la bande cubitale est réduite 4 un
mince liseré longeant le bord de la cellule et envoyant peut-étre des pro-
longements le long de la base des nervures médianes et cubitales, comme
chez lhecuba actuel. La bande radiale est étroite et trés fine ou absente a
la base et la bande anale étroite et trés courte. La coloration est uniforme-
ment verte.

Le dessus des ailes postérieures est toujours largement jaune translucide
dans sa partie antérieure, avec quelques taches submarginales concolores
et 5 taches discales noires. La couleur métallique est verte, mais la cellule
I est noire et la base de laile peut étre sablée de noir.

Les dessins du dessous des ailes sont probablement fort semblables a
ceux de la 2 de lydius. Les antérieures montrent une bordure marginale
noire assez large et une série de taches discales confluentes en une ligne
assez reguliere au milieu des cellules 2 4 8. Aux ailes postérieures, le bord
est egalement margé de noir, les taches discales grandes, rondes et au
nombre de six, le tornus teinté de jaune et peut-étre de noir et la cellule
et les nervures médianes et cubitales bordées de noir. L’ensemble des deux
ailes nest pas métallisé.

La cellule des ailes antérieures est relativement large, cest-a-dire en-
viron 2,5 fois moins large que longue alors qu’aux ailes postérieures, elle
est au moins 3,5 plus longue que large.

Les valves sont légérement ovales, la harpe assez gréle et bien allongée,
avec un col et une téte faiblement indiqués et des épines en position
apicale. Il y a deux dents basales assez longuement triangulaires.

Cet ancétre parait avoir donné naissance a deux formes représentant le
début de deux directions d’évolution différentes: la lignée de croesus con-
tenant ce dernier, aesacus et priamus et la lignée @allottei, comprenant en
outre celui-ci, victoriae et alexandrae.

L’ancétre théorique 2 (fig. 9), progéniteur de la lignée de croesus a as-
sez peu changé extérieurement, mais acquis de nombreuses spécialisations
de détail. Le mésonotum est orné d'une bande métallique verte longitudi-
nale. Aux ailes antérieures, la bande radiale s'est un peu renforcée: elle
est étroite mais bien réguliere des sa base. La bande cubitale est
présente mais linéaire et la bande anale toujours courte. Les ailes pos-
térieures sont comme celles de lancétre I, avec Taire jaune translucide
réduite et localisée 4 la moitié antérieure de l’aile, mais elles ont acquis une

98

priamus

rouge présente,

aesacus

croesus

cell. ant. étroite,
cell. post. courte.

Bande anale longue,

couleur dorée.
aire transluc. perdue.

Bande mésonotum présente, 2
bande radiale réguliére, es
bande anale courte,

dire transluc, réduite,

en position antérieure,

cellule | post. métallique,

JOURNAL OF

Pilosité thoracique

cell. | post. métallique,
deux dents 4 la harpe.

Bande radiale large,
dire transluc. conservée,

THE LEPIDOPTERISTS SOCIETY

victoriae

alexandrae

allottei

28 caractéres de plus
en plus accusés

3, Bord post-oculaire perdu,
pilosité thorac. rouge présente,
ailes allongées, arrondies, bicolores,
bande radiale mince a la base,
bande anale tres courte,
zone translucide perdue,
cellule | post. noire,
valves ovales, harpe avec
téte et col bien marqués.

valves subdiscoidales, 1
4 A
harpe en parallélogramme, “\ Bord post-oculaire présent,
.deux dents basales réduites. Bille THenae, couse p<ecnne
: :

Bande radiale présente,
nines

bande cubitale en large

triangle,

Schoenbergia
pas de croissant androconial,

post. largement translucides.

#

Troides

Trogonoptera

Ailes antérieures”
lignées de blanc.

Ailes antérieures avec”
série de triangles verts.

Fig. 9.

Ornithoptera

ailes triangulaires, vertes, unicolores,
bande radiale mince a la base,
bande cubitale linéaire,

bande anale courte,

croissant androconial présent,
aire translucide présente,
cellule ant. large,

cellule post. longue,

valves ovales, harpe avec
téte et col un peu marqués,
deux dents basales.

Arbre phylétique du sous-genre Ornithoptera. La divergence relative des

tiges indique les directions différencielles d’évolution. Les niveaux ot se situent les
especes signifient trés approximativement les degrés relatifs de spécialisation de ces

derniéres.

VoLUME 24, NuMBER 2 99

importante spécialisation, la cellule I est entierement métallique en-dessus.
La nervulation reste inchangée, avec la cellule des ailes antérieures assez
large et celle des postérieures plutot courte. Les génitalia, par contre, sont
assez modifiés: les valves sont subcirculaires, la harpe courte, assez massive,
en forme de parallelogramme, avec un col et une téte 4 peine marqués. Les
deux épines basales sont trés peu proéminentes.

Cet ancétre 2 a donné naissance aux trois especes actuelles croesus,
aesacus et priamus, occupant des seuils de spécialisation fort inégaux.
Croesus na acquis que peu de particularités nouvelles, mais ces dernieres
sont telles que aspect de Tinsecte a beaucoup change. Les ailes anté-
rieures se sont étirées vers l’extérieur en triangles assez obliques. La bande
radiale sest considérablement élargie, la bande cubitale a enticrement dis-
paru, alors que la bande anale a conservé son aspect primitif et reste
courte, sauf chez certains individus chez qui elle se prolonge en une
trainee sablée jusqu’a M1 (lydius berchmansii vd. Bergh). Le caractere le
plus frappant est que la lumiére interférencielle primitivement bleue
des aires métalliques, qui se combine avec le pigment jaune sous-jacent
pour donner une couleur apparante verte, a viré au rouge et produit ce
flamboiement orange qui a tant ému Wallace et valu son nom a lespece.
Par conséquence, abdomen qui était primitivement jaune, a aussi viré a
Yorange. Seule la bande du mésonotum reste verte.

Priamus est considérablement spécialisé et a acquis une série de parti-
cularités qui pourraient faire figure de caractéres primitifs comparés a
ceux dallottei et de victoriae, mais qui sont en réalité des spécialisations
dans une direction d’évolution opposée. Les zones noires de la base de
labdomen, des pleurites, du VIIle tergite et des valves sont nettement
réduites. Les ailes antérieures sont en triangles rectangles, caractere
primitif, mais le bord apical des postérieures est fortement crénelé. Sur
le dessus, les bandes radiale et cubitale sont inchangées, mais la bande
anale se prolonge de fagon continue tout le long du bord anal, jusqu’a
l'apex de laile devenant remarquablement symétrique de la bande radiale,
specialisation importante qu’on ne trouve que chez priamus et aesacus.
Les ailes postérieures ont perdu la large zone jaune translucide, mais en
ont retenu des traces, inconstamment présentes, sous forme de petites taches
submarginales, dans l’aire sous-costale et les cellules III a VI. Les spéci-
mens possédant ces taches ont recu des noms variés: triton Felder chez
poseidon, flavomaculata Przegendza et Ribbe chez hecuba et urvilleanus
respectivement et flavopunctata Rousseau-Decelle chez caelestis. La base
de laile postérieure a tendance a se sabler de noir. Les dessins du dessous
des ailes sont inchangés, de méme que les génitalia. Enfin, la cellule des

100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ailes antérieures est nettement rétrécie et celle des postérieures raccourcie,
spécialisation exclusive a priamus.

La différenciation des nombreuses sous-especes de priamus a ete bien
étudiée par Zeuner (p. 138, 148). Je n’y reviens donc pas ici.

I] nvest difficile d’assigner une place précise a aesacus, car il nvest resté
inconnu. Je puise donc les renseignements qui suivent chez Zeuner (p.
137, 149). Aesacus s’est probablement différencié dune forme voisine de
croesus et occupe un seuil de spécialisation moins élevé que priamus, ce
quindiquent clairement la cellule des ailes antérieures restée large et
celle des postérieures toujours longue, comme chez croesus. La bande
radiale des antérieures est large quoique moins que chez croeésus et la
bande anale tres longue comme chez priamus. Aux ailes postérieures, la
cellule I est métallisée. La coloration générale est bleu verte. Les valves
sont subcirculaires et les dents basales de la harpe sont réduites, mais celle-
ci est amincie a son extrémité et avec une armature spineuse apicale sub-
circulaire comme dans la lignée d’allottei.

L’ancetre théorique 3 de la lignée d‘allottei (Fig. 9) montre peu de
specialisations fondamentales a partir de Yancétre I, mais de nombreux
changements de detail. Les bandes post-oculaires blanches sont perdues,
mais la pilosité rouge des pleures thoraciques est conservée. L’abdomen
est toujours jaune vif avec les zones noires basales et latérales bien dé-
veloppées et sur les segments 3 et 4 apparait une ligne décolorée. Les ailes
antérieures sont en triangles obliques comme chez croesus, avec leur angle
apical postérieur arrondi. Les postérieures sallongent légérement et
léchancrure de la cellule I s'approfondit. La coloration des antérieures est
primitive, avec la bande radiale trés mince a la base de Ilaile, la bande
cubitale toujours présente mais vestigiale et la bande anale courte. Aux
ailes postérieures, la coloration du dessus a changé dans le méme sens que
celle de priamus et ressemble beaucoup a celle de ce dernier. La zone
basale, jaune translucide a entiérement disparu, mais les point submargi-
naux de la méme couleur sont retenus. La cellule I est toujours noire. Aux
deux ailes, la coloration bipartite apparait: a la partie apicale, le vert se
teinte nettement de jaune. En dessous, la bordure marginale noire se
réduit, mais le noir a tendance a remonter l’extrémité des nervures. Les
taches discales sont légérement réduites et la couleur de fond se métallise.
La nervulation est primitive, de méme que les génitalia.

Allotiei montre des changements assez nombreux mais légers A partir de
ancctre 3. Il est plus proche de ce dernier que croesus lest de son ancétre
2. La pilosité thoracique rouge est maintenant perdue. Les ailes antérieures

ont retrecies et arrondies, alors que les postérieures se sont un peu
| ees. La bande radiale des ailes antérieures est évanescente sur toute

mvaa

VoLUME 24, NUMBER 2 101

sa moitié basale, comme cest parallélement les cas chez chimaera et
tithonus. La bande anale est nettement élargie et se prolonge en un sable
jusquau bord costal. Les taches discales des ailes postérieures sont ré-
duites en nombre et de taille. Mais surtout la couleur métallique du des-
sus des deux ailes est maintenant nettement bipartite: aux antérieures,
Tapex de la bande radiale et aux postérieures le pourtour de laile sont
vert-jaune assez brillant. Sous les deux ailes, les dessins sont toujours
dun type nettement primitif, sauf que le long du bord des postérieures le
noir forme de nets triangles a l'apex des nervures et la couleur générale se
métallise. Aux antérieures, la cellule a continué de s’élargir alors quelle
s est encore allongée aux postérieures. Les génitalia sont peu changes sauf
qu une des dents basales de la harpe a disparu, alors que l'autre sest un
peu allongée.

Victoriae est une espéce tres remarquable, qui représente l’aboutisse-
ment de lévolution de la lignée dallottei, et, comme telle, occupe un
palier de spécialisation nettement plus élevé que priamus. Elle apparait
tres spécialisée par tous ses caractéres et cela dans la méme direction
dévolution quwallottei. Ceci indique quelle est issue soit de ce dernier
soit d'un ancétre extrémement semblable a celui-ci. Victoriae et allottei
représentent donc deux étapes d'une spécialisation linéaire. J’ai noté non
moins de 28 caracteres qui sont moyennement spécialisés chez le premier
et tres accentués chez le second. Ce sont: 6, 7, 10, 11, 12, 13, 14, 15, 17,
om Oelhe 245 27, 23529) 30) 315 32, 34, 35, 36, 37, 38, 39, 40 et Al.
Comme ils sont cités dans la table qui précéde, je ne les énumérerai pas
une seconde fois, mais commenterai un petit nombre des plus intéressants.
Les plus frappants sont évidemment l’élongation et l’arrondissement de la
forme des deux ailes et ’exagération des proportions des deux cellules. Le
plus intéressant est certainement la forte modification du développement
des bandes métalliques des ailes antérieures. La bande radiale nest pas
interrompue en son milieu comme il pourrait le sembler et comme Zeuner
Ya cru (p. 139). Mais une comparaison avec allottei et les divers Schoen-
bergia montre que cette bande s’est retirée, comme en une sorte de mouve-
ment amoebien, de presque toute la longueur de laile et quelle s est
groupee et considérablement élargie a son extrémité pour former un tri-
angle doré remplissant lintérieur de la partie apicale de Taile. La bande
anale sest considérablement développée, non en longueur comme chez
priamus, mais en largeur et occupe toute la base de Jaile, atteignant
[aire radiale ou elle a pris la place de la bande de ce nom. La bande cubit-
ale, quia été perdue par allottei déja, n'est pas comprise dans cette large
aire basale.

Notons aussi que les taches submarginales jaunes translucides, jamais

102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

signalées chez allottei, sont presque toujours présentes chez victoriae et
atteignent parfois une taille considérable. Si, chez allottei, tout le pour-
tour de laile postérieure est vert jaune, chez victoriae, seul lapex est doré
et il peut étre tres brillant.

Certaines des variations intra-subspécifiques de victoriae sont inte-
ressantes, car elles se font dans une direction opposée a celle de leurs homo-
logues de priamus. Chez ce dernier, les taches discales noires et les points
jaunes submarginaux du dessus des ailes postérieures décroissent de taille
d’avant en arri€re et, lorsque le centre de laile est poudré de noir, cest a
Ja base de laile quil lest le plus fortement. Chez victoriae, le gradient
d’évanescence a renversé sa direction: cest pres du tornus que les taches
noires et jaunes sont les plus développées et elles décroissent en direction
de lavant. De méme, lorsque la surface sennuage de noir, cest a partir
de la bande dorée apicale.

Je ne crois pas inutile d’ajouter quelques commentaires sur les différen-
tes sous-especes et aberrations de victoriae. Les sous-espéces décrites
jusqu ici sont les suivantes:

regis Rothschild, de Bougainville.

isabellae Rothschild, de Santa Isabel.

reginae Salvin, de Malaita

victoriae Gray, de Guadalcanal, Tulagi et Florida.
rubianus Rothschild, de Rubiana et Kolombangara.
resplendens Ehrmann, de Choiseul.

La description d'une septieme sous-espece réecemment découverte a San
Cristobal est annoncée.

A mon avis, resplendens est certainement synonyme de regis comme
cest probablement le cas disabellae. Des quatre espéces bien connues
regis et victoriae sont certainement les plus primitives, ce qu indiquent le
maximum d’extension de la couleur métallique sur le dessus des deux ailes
et les deux couleurs vert et or bien contrastées. Victoriae toutefois montre
une réduction en longueur de la bande radiale, ce qui est une légére
specialisation par rapport a regis. Ces deux sous-espéces sont extréme-
ment variables, surtout regis et cela aussi bien par les couleurs que par les
les dessins. J'ai vu des spécimens dont le dessus des deux ailes est dun
vert uniforme, acide et agressif. Chez certains autres, la bande apicale des
ailes postérieures a le brillant lumineux de Yor en fusion ou l’éclat adouci
du vieil or patiné ou méme la nuance rougeatre du cuivre oxydé. Chez
quelques exemplaires, le centre des ailes postérieures peut étre
]

dun bleu ciel trés clair. De nombreux spécimens de chacun des

deux regis et victoriae présentent les caractéres de Yautre, ce qui
f i (111Ee@ fac dar =| pod rpm 7 \ . OW ° .
lit qi es dernieres ne sont guere identifiables que statistiquement.

VoLUME 24, NuMBER 2 103

I] nest pas impossible que regis aussi ne soit pas valide et ne soit qu'une
simple race de victoriae. Reginae est nettement plus spécialisé que les
deux précédents, par ses ailes postérieures largement envahies de noir a
partir de la bande dorée, qui est elle-méme réduite. Rubianus enfin est la
forme de fin de lignée et la plus modifiée par sa coloration appauvrie. La
bande radiale des ailes antérieures est fortement réduite de taille dans sa
partie antérieure, ce qui fait quelle se trouve maintenant au milieu de la
largeur de Vaile et elle est retournée au vert primitif. La bande apicale
dorée des ailes postérieures de méme que les points jaunes submarginaux
sont perdus.

I] est intéressant de noter que, comme chez priamus, les mémes varia-
tions sont visibles chez chacune des sous-espeéces: la couleur noire peut
envahir plus ou moins largement les ailes postérieures, le vert et lor sont
plus ou moins contrastés et les deux bandes des ailes antérieures peuvent
étre réunies par un sablé métallique.

Un certain nombre de formes individuelles ont été décrites. Buinensis
Le Moult et infanta Bryk et Peebless désignent tous deux des regis dont les
deux bandes des ailes antérieures sont réunies. Sanguinea Rousseau-De-
celle, brabanti Le Moult et gabrielli Le Moult s’appliquent a des regis
dont certaines régions des deux ailes, en général les aires dorées, portent
“une surcharge rouge-brun, couleur de sang séché”. Cette couleur a
également été signalée chez chimaera draceana ab. sanguifluens Rous-
seau-Decelle. J'ai moi-méme vu des regis pourvus de zones rougeatres,
variables en intensité et en extension, parfois réduites a de petites taches
irrégulieres et asymétriques aux deux ailes. Ces irrégularités pourraient
indiquer une malformation ou une affection pathologique des écailles
meétalliques. Alexisi Le Moult désigne des regis dont le vert a entierement
tourné au doré sur les deux faces des deux ailes. Lanieli Le Moult, enfin,
est intéressant car il définit des regis qui ont retenu sur ou sous les ailes
postérieures de deux a quatre taches discales noires, homologues de celles
de priamus et allottei.

Reste a placer la troisieme et derniere espece de la lignée dallottei,
alexandrae. Cette forme étant célébre et ses caracteres bien connus, il me
parait inutile de m’y attarder longuement. Elle est vraisemblablement is-
sue de lancétre 3 et non d'une forme voisine d’allottei, car elle a conservé
quelques caractéres primitifs perdus par ce dernier, comme la pilosité
thoracique rouge, une bande cubitale vestigiale et les deux dents de la
base de la harpe. Elle posséde aussi quelques particularités en commun
avec allottei, comme la bande radiale des ailes antérieures nuageuse a sa
base et vert-jaune a son extrémité, les proportions des cellules des deux
ailes et ’abdomen resté jaune vif. Avec victoriae elle montre en commun

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

croesus
priamus
aesacus
allottei
alexandrae
victoriae

Fig. 10. Arbre phylétique du sous-genre Ornithoptera d’aprés Zeuner.

la forme longuement elliptique des ailes et la courte frange anale blan-
chatre. Mais alexandrae a développé un bon nombre de caracteres de son
cru et des plus importants, ce qui montre quelle s'est tres fortement dif-
férenciée de lancétre 3 dans une direction particuliére. Citons le gigan-
tisme, la perte presque complete du pigment jaune sur les deux ailes, la
bande anale élargie, tres allongée et englobant le croissant androconial,
les dessins tres particuliers des ailes postérieures, dont la cellule I est
colorée comme dans la lignée de priamus et la perte de toutes les taches
discales sous les deux ailes. Le dessous des ailes est trés largement coloré
et lune des dents basales de la harpe est barbelée, caractéres acquis par
convergence avec les Schoenbergia.

Nous référant a nouveau a larbre phylétique de Zeuner (Fig. 10), nous
voyons maintenant les changements qui s'imposent. Cet auteur a reconnu
l'existence des deux lignées du sous-genre, celle de priamus et celle Wal-
lottei. Pourtant, il indique que croesus s’est différencié le premier du tronc
de Tarbre, avant Yapparition des deux lignées. Ensuite seraient apparu
simultanément priamus, aesacus et Yancétre de la lignée dallottei qui a
son tour aurait produit simultanément allottei, alexandrae et victoriae.

In realité, il semble assez évident que lancétre du sous-genre a donné
naissance d’abord aux deux ancétres 2 et 3, A partir desquels se sont dif-
férenciées les deux lignées. Croesus semble en effet étre apparu le premier
dans la lignée de priamus, mais dans celle d’allottei, il est probable quale-
xandrae se soit différencié avant allottei.

VoLUME 24, NuMBER 2 105

Bibliographie

RoruscHiLp, W. 1914. Description of a New Troides. Nov. Zool. 21:275.

NiEPELT, W. 1916a. Lepidoptera Niepeltiana 2, p. 20-21, pl. 17, fig. 1-2.

NrepeLt, W. 1916b. Beitrag zur Kenntnis der Ornithoptera alottei Rothschild. Ent.
Rundsch. 33:31.

PrEEBLESS, H. M., W. ScHMAsSsSMAN. 1917. Description of the female of Troides al-
lottei Rothsch. Nov. Zool. 24:426—427.

ZEUNER, F. FE. 1943. Studies in the Systematics of Troides MWHiibner and its Allies.
Trans. Zool. Soc. 25. (3), 184 pp.

STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN
NEW ENGLAND. I. ABUNDANCE AND SEASONAL OCCURRENCE
OF THE SPECIES, 1961-1969

THEODORE D. SARGENT
Department of Zoology, University of Massachusetts, Amherst, Massachusetts

and

SIDNEY A. HESSEL

Entomology Section, Peabody Museum, Yale University, New Haven, Connecticut

The Catocala moths pose an interesting evolutionary enigma. This
large genus, consisting of some 200 species in the broadest use of the
generic name, is characterized by extensive sympatry throughout most
North Temperate regions of the world, and its members occupy a rela-
tively narrow ecological niche, the larvae being oligophagous and the
adults being cryptically marked when at rest on tree trunks. In New
England alone, at least 52 Catocala species are known to occur (Forbes
1954), and we have taken as many as 21 species on a single night at one
location.t Nevertheless, natural hybrids are unknown. This tremendous
array of sympatric and rather closely related species immediately raises
the question as to what sorts of mechanisms operate to limit natural
hybridization within the genus. One long-range goal of these studies is to
describe such mechanisms through detailed analyses of the behavior,
ecology, and genetics of these moths.

Despite the popularity of the North American Catocala moths with
collectors, relatively little of their total biology is known. They have
been treated taxonomically by Grote (1872, 1873, 1876), Hulst (1880,

120 August 1961, Washington, Connecticut: C. epione, antinympha, badia, habilis, flebilis, ob-
scura, residua, retecta, dejecta, palaeogama, subnata, neogama, ilia, parta, concumbens, sordida,
andromedae, ultronia, grynea, praeclara, and amica.

106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1884), Hampson (1910), McDunnough (1938), and Forbes (1954).
Plates depicting most of the North American species are available in
Holland (1903) and Barnes and McDunnough (1918). Present know]-
edge of the life histories and geographic ranges of the New England
Catocala is summarized in Forbes (1954).

Information on the behavior and ecology of these moths is scanty and
largely anecdotal. Some observations on the resting habits of adults have
been recorded (e. g. Bunker, 1874; French, 1880; Johnson, 1882; Rowley
and Berry, 1909; Kettlewell, 1958; Sargent and Keiper, 1969). Limited
data on the movements of adults have been obtained from studies of
color-marked individuals (Brower, 1930). Virtually nothing is known
of courtship and mating behaviors—a fact that has precluded any detailed
studies on the genetics of these moths.

The present report constitutes a first step in our study of the Catocala
of southern New England: an account of the species that are present,
based on daily counts of adults taken for several years in two localities.
It is hoped that these data will provide a foundation for planning further
investigations, will serve as a record for future comparative purposes,
and will stimulate others to obtain comparable data from their localities.

Methods

The basic data utilized in this report are daily counts of adult Catocala
taken in the vicinity of Amherst, Massachusetts (Area 1) for the years
1964 through 1969; and from Washington, Connecticut (Area 2) for the
years 1961 through 1965, 1967, and 1969.

Area I. Records here are based on collecting of 2009 specimens by T.
D. Sargent and two graduate students at several localities in Franklin
and Hampshire Counties in north-central Massachusetts, all localities lying
within 10 miles of the center of Amherst. The vast majority of the records
are from two sites, one in Pelham (1964-66) and another in Leverett
(1967-69). Catocala taken at bait, a brown sugar-beer mixture painted
onto tree trunks, comprise 84% of the records from this area, whereas
Catocala taken at lights (several 150-watt incandescent bulbs, and one
L5-watt black light fluorescent tube) comprise 12% of the records. The
additional 4% of the records are based on the finding of resting moths in
nature, Catocala were recorded on a daily basis from 1 July to 1 Septem-
ber (except for occasional 1 to 2 day absences, or days of extremely inclem-
ent weather), and more sporadically from 1 September to 1 November
each Year,

Area 2, Records here are based on collecting of 5806 specimens by S.
A. Hessel at one site in Washington, Litchfield County, in west-central

VoLUME 24, NuMBER 2 LOT

Connecticut; this site lies approximately 65 air-miles to the southwest of
Amherst, Massachusetts.? All records of Catocala from this area are based
on specimens taken at lights, most at one Robinson mercury vapor light-
trap, and some at one 15-watt black light fluorescent tube. These lights
were in operation from mid-March to mid-November each year (except
for occasional 1 to 3 day absences ).

Several important differences in collecting procedures between the
two areas should be stressed:

(1) Captures in Area 1 were predominantly at bait; captures in Area
2 were exclusively at lights.

(2) Collecting in Area 1 was generally terminated by midnight; in
Area 2 the Robinson trap was operating continuously from dusk to dawn.

(3) Several collecting sites are included in Area 1; Area 2 includes
only one collecting site.

(4) Collecting in Area 1 was sporadic after September 1, but con-
tinued unabated in Area 2.

Because of these differences, the records for the two areas will be pre-
sented separately.

It should also be noted that some Catocala individuals may have been
recorded on more than one occasion, as the majority of specimens in both
areas were released after examination. However, studies of color-marked
Catocala in Area | (Sargent, in prep.) indicate that very few individuals
are captured on more than one occasion.

The species of Catocala were identified as keyed and described in
Forbes (1954), and the species names used throughout this report are from
that source. It should finally be noted that specific distinctions were not
always made between gracilis and sordida in both areas, and among
crataegi, blandula, and mira in Area 1. It is known from mounted speci-
mens, however, that all of these species occurred in both areas.

In area 1, the sex of Catocala specimens was determined, and the
precise time of their capture noted. These data are not included in the
present report, but will be treated in subsequent papers.

Results and Discussion

Abundance. The Catocala taken each year in Areas 1 and 2 are enumer-

2 This site lies at the bottom of a very narrow north-south valley through which an all-season
stream flows southward. The lower end of the valley, only one mile distant, opens onto terrain which
rolls gently southward to the Connecticut coast, while northward, the Litchfield Hills, of which it is
part, become the Berkshires of Massachusetts. The surrounding hills are largely mixed deciduous
woodlands, but include several seral stages resulting from the abandonment of farms, pastures, and
woodlots of the Colonial period. Climatically, as demonstrated by meteorological maps, the site is to
be included with territory considerably more northemm than with the Connecticut and Hudson River
valleys of its latitude.

108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Catocala spp. recorded in the Amherst, Mass. area, 1964—69.

Totals and
Numbers of Individuals Percent of
& (Percent of Yearly Total for All Species ) Grand Total
—=. stouseaatine

Species 1964 1965 1966 1967 1968 1969 Period
ilia 12 49 20 AT 206 346 680
(22) (25) Ome) Mes (45) (34)

ultronia 7 49 ILS 25 49 79 324
(3). (5) - G2) @) Cae (16)

crataegi et al.* 2), 14 AT 5 36 5 109
(4) (7) (16) (1) (10) (1) (5)

retecta = 3 iu 30 D 58 107
(2) (4) (8) (1) (8) (5)

antinympha 1 I alts) ih 11 19 104
@ @ 6) Ge Grams (5)

gracilis & 6 2 LiL ih 5 10 91
sordida (hE) (1) (4) (16) (iL) (ab) (5)
cara i _ 13 2 3 60 79
(2) (4) (i) (1) (8) (4)

concumbens 3 5 19 14 6 29 76
(5) (3) (6) (@) (2) (4) (4)

relicta — DAIL 5) iLil 3 Iti 51
(11) (2) (3) (1) (1) (3)

grynea 2 18 10 i 10 50
(4) (9) (3) (2) (1) (1) (2)
amica 3 4 4 30 3 6 50
(5) (2) (1) (8) (i) (1) (2)
amatrix _ — — _ = 36 36
: (5) (2)
unijuga 3 3 5 6 6 10 Bie)
| ® @ © @- Cae (2)
micronympha 2 4 2 22, — 3 33
(4) (2) (1) (6) (2)
epione il il - 6 8 16 32
(2) (1) (2) (2) (2) (2)

andromedae 2 py) 8) 15 4 4 30
(4) (1) (1) (4) (1) (1) (1)

praeclara 5 if 6 co) 2 29
. (9) (4) (2) (1) (1) (1) (1)
similis A 6 _ LiL E ed pal
(7) (3) (3) (1)

cerogama ue a 9 3 il 7 ite
fox (1) (1) (1) (1)
residua ~ _ —_ = — 11 Gah

1

palaeogama ae = 1 2! 1 e
Cay (iy)

neogama -- ne 1 re} il 5 10

(1) (1)

VoLUME 24, NuMBER 2 109

TABLE 1. Continued.

Totals and

Numbers of Individuals Percent of
& (Percent of Yearly Total for All Species ) Grand Total
for Entire
Species 1964 1965 1966 1967 1968 1969 Period
habilis — - — 5 — 3 8
(1)
coccinata | — I, — 5 1 1 8
(1) (1)
obscura I 3 il - _ Il 6
(2) (2)
briseis — 1 2 - _ — 3
(1) (1)
innubens - i - — - - iL
(1)
piatrix — — iL — — - iL
dejecta — — — — - IL 1
parta — = = = = iL
No. Species 16 20 Dll al IL 26 ao
No. Individuals 55 195 294 366 354 745 2009

* records of blandula and mira included here

ated in Tables 1 and 2 respectively. In both cases, the species are listed
in a descending order of overall abundance. Examination of these tables
reveals some differences between results from the two areas. Catocala
numbers, in terms of both species and individuals, are greater from Area
2. In part, this difference must be due to the longer daily and seasonal
collecting periods in Area 2, and perhaps to a greater efficiency of the
Robinson trap, when compared to bait, as a collecting device. However,
further consideration indicates that those species that are markedly more
abundant in Area 2 are almost invariably hickory and walnut (Juglan-
daceae) feeders. Accordingly, the data were reanalysed with reference
to the known foodplants of the various Catocala species (Forbes, 1954),
and this procedure revealed that remarkably more hickory-walnut feeders,
both in terms of species and individuals, were taken in Area 2 (Table 3).
This disparity in records of feeders on the Juglandaceae seems to reflect
something more than the previously listed differences in collecting proce-
dures between the two areas. It seems more likely that differences in the
frequency of hickories and walnuts are important. In this regard, the
presence of Butternut (Juglans cinerea L.) as a common tree in the wood-
lands near the collecting site in Washington, and its virtual absence in all
collecting areas near Amherst, seems most suggestive. It could be that

110 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 2. Catocala spp. recorded in Washington, Conn., 1961-65, —67, -69.

Totals and
Percent of
Numbers of Individuals Grand
& (Percent of Yearly Total for All Species ) Hoo
Species 1961 1962 1963 1964 1965 1967 #1969 Period
palaeogama 194 487 ILD, 40 85 81 48 895
(15) (34) (4) (8) (6) @) (8) (15)
residua 216 345 45 53 5A 135 39 887
Cm) (4) Cle) ID) a) (7) (15)
habilis IEPA 79 63 119 78 159 34 653
(9) (Ss) Gi) (4) G4) G4 (6) (11)
amica 85 78 10 Di, Dil 89 50 366
(7) (6) (3) (9) (5) (8) (9) (6)
antinympha By) 26 13 13 16 Ie aT 329
(4) (2) (4) (2) ©) ©) (5) (6)
neogama 97 36 8) 38 39 68 28 329
(8) (3) (8) (7) (7) (6) (5) (6)
concumbens 14 1) Dy 59 68 Ve 60 309
(1) (1) () Cat) de) (6) (10) (6)
retecta 83 5S) 19 29 34 55 19 294
(7) @ (6) (5) (6) (9) (3) (5)
grynea 2 30 8 19 35 39 56 259
(6) (2) (3) (4) (6) (3) > 1G) (4)
ultronia 4] 38 14 24 32 45 OM eh
(3) (3) (9) (5) (6) (4) (6) (4)
epione 69 ot 14 6 19 24 15 184
(9) (3) (9) @ (3) (2) (3) (3)
andromedae AQ 30 7 8 26 10 18 139
(3) (2) (2) 5) (5) (1) (3) (2)
obscura 27 17/ 3 17 9 26 95 124
(2) (1) (1) (3) (2) (2) (4) (2)
micronympha 38 43 5 11 4 6 ile 122
8 @ @ @ @ @ eae
serena 3 ~ = 8 1D 52 28 103
() (2) (5) (5) (2)
gracilis & 16 “il 12 15 10 7 8 Gls;
sordida (1) (4) (3) @ (1) (1) (1)
mira 4 14 § Uh 9 Y 20 68
(ly) (3) (2) (2) (3) Gy
hadia 9 7 3 2 § DL 7 57
(1) (1) Q @ Gi
cara 15 1 B22 3 6 Y) 5 4 AT
| (1) (1) (1) (1) (1) (1)
dejecta 0 "i ] 2 5) 15) IL 39

September 1956, one specimen.

VoLUME 24, NuMBER 2 aT
TABLE 2. Continued.
Totals and
Percent of
Numbers of Individuals Grand
& (Percent of Yearly Total for All Species ) Total. for
SS ntire
Species NOG 962 1963) = 19645 1965) 196i 1969!) Renod
ilia 9 4 il 5 9 4 5 37
(1) (1) (2) (1) (1)
judith i = — = il 9 94 35
OO @
subnata 9 19 2 2 1 i - 34
(1) (1) @ (1)
parta 9 2, 6 3 4 0 2 33
(1) (2) (1) (1) (1) (1)
unijuga ih 5 3 6 4 8 — 33
@) (1) (1) (1) (1) (1)
similis a il 1 - il 4 - 20
(1) |
flebilis 10 2) iL = 1 4 iL 19
(1)
coccinata 7 4 iL 2) 1 Y 2 19
(1)
praeclara it - i 1 4 6 4 17
@& (1) @
relicta 6 = 1 1 5 2 ] 16
(1)
blandula = 6 3 1 2, 2 - 14
(1)
crataegi vel 2 IL i ~ 2, 1 8
innubens i = = — D TL = 4
amatrix — 1 iL - — — - 2
- briseis = = = = = 2 = S)
piatrix Jk _ - - - - - i
cerogama — - - - _ 1 = il
vidua = = = iL = = = 1
No. Species oo 29 31 30 Bo) 35 28 Soe
No. Individuals 1275 1412 306 530 ies LING yIL 579 5806
* During the period 1952-60, one additional species was noted in this area—C. robinsonii, 15

112 JouURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 3. Foodplants of the Catocala spp. from two localities.

Amherst, Mass. Washington, Conn.

No. % Total No. % Total

Foodplants Species Individuals Species Individuals
Salicaceae? i 13.9 7 7.6

Willow (Salix);
Poplar (Populus)

Myricaceae” 1 5.2 2 6H
Bayberry (Myrica);
Sweet Fern (Comptonia)

Juglandaceae® 9 9.3 14 62.0
Walnut (Juglans);
Hickory (Carya)

Fagaceae* 5 39.4 5 9.7
Oak (Quercus)
Rosaceae? 5 24.0 5 10.0

Apple (Pyrus);
Thorn (Crataegus);
Cherry (Prunus )
Ericaceae® 3 6.0 8) ond
Blueberry (Vaccinium );
Andromeda (Andromeda)
Others‘ 3 all 3 0.4

Catocala species included:

1 yelicta, parta, briseis, unijuga, cara, concumbens, amatrix

2 antinympha, badia

2 piatrix, epione, habilis, serena, judith, flebilis, obscura, residua, retecta, dejecta, vidua, palaeo-
gama, subnata, neogama

4 ilia, coccinata, similis, micronympha, amica

° ultronia, crataegi, mira, blandula, grynea

6 gracilis, sordida, andromedae

7 innubens (Gleditsia), cerogama (Tilia), praeclara (Quercus ?, Crataegus ?).

Area | is near or beyond the northern limit of some of these Catocala
species, due perhaps to a sparse distribution of certain foodplants, or to
climatic marginality for the insects themselves.

The presence and absence of a foodplant seems to explain the single
remaining difference in species between Areas 1 and 2 (i.e. bayberry,
Myrica pensylvanica Loisel, the foodplant of C. badia, is present only in
Area 2).

[t is also possible, of course, that some differences in records from the
two areas are due to differences in collecting procedures, which reflect,
in turn, behavioral differences among the Catocala species. For example,
C. ilia has been taken much more often at bait than at lights (96% of 371
captures in Leverett, 1967-69), and this fact may well explain its ap-

rently higher numbers in Area 1] (and the consequent higher percentage
i feeders on the Fagaceae). In contrast, C. antinympha has been more

1
‘

's than at bait (88% of 74 captures in Leverett, 1967-69),

VoLUME 24, NuMBER 2 ERS

TABLE 4. Early occurrence of several common Catocala species in two localities.

Extreme Early Date Median Early Date
Month/Day Month/Day
Species Amherst Washington Amherst Washington
antinympha rials yale 7/22 Ty Mel
retecta 8/2 7/30 8/11 8/10
concumbens . UP2A M23 THB THe
ultronia Hef Malt Ws 7/20 7/20
grynea 7/14 TMP 1/22 1p PAL
amica 1/2, ee Tf LS ome

and this could account for its apparently higher frequency in Area 2. Dif-
ferences of this sort, as well as differences in the time of flight of the
various species during the night, are being carefully studied, and will
form the basis of further papers in this series.

Changes in Abundance. The records presented here do not cover a
sufficient number of years to allow extended analyses of changes in
abundance. However, the variations and trends in annual numbers of
certain species warrant brief comment.

Several species exhibited wide fluctuations in annual abundance (e.g.
C. ilia and C. ultronia in Area 1, C. palaeogama and C. habilis in Area 2,
and C. antinympha in both areas). Two species in Area 1, C. amatrix and
C. residua, were never recorded until 1969, and then both were relatively
common. These fluctuations indicate that detecting long-term trends in
Catocala abundance may often require longer series of annual records
than those considered here.

The records from Area 2, together with notes of S. A. Hessel extending
back to 1952, do indicate that two species, C. serena and C. judith, have
recently become remarkably more common in that area. Although neither
of these species was collected in Area 2 prior to 1960, since then 104
serena and 35 judith have been collected. These results could be due to
erratic long-term oscillations in the abundance of these species, or might
represent their recent range extension from more southern regions into
Area 2. If these two species are presently extending their ranges, then we
might eventually expect to take specimens in Area 1, where neither has
yet been taken.

In all of the records from both areas, only one species, C. subnata,
seems to be showing any evidence of a recent decline in numbers. This
species is known as one which exhibits long-term fluctuations in abun-

dance (Forbes, 1954).

114 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

TABLE 5. Seasonal occurrence of Catocala.

Percent Occurrence

July August September October
Species! —____ eee
(N) 1-15 16-31 1-15 16-31 1-15 16-30 1-15 16-31
blandula 61 28 6 6
(18)
coccinata 30 30 33 7
(27)
similis 12 68 7 2
(41)
micronympha NB AT 22, 5 Il
(155)
ilia il AT 35 12 4
(fale)
ultronia 1 Al 39 7 1 -
(555)
mira 3 Al 42 14
(69)
epione 2, on 46 15
(216)
andromedae 2 On 39 Pall 1
(169)
antinympha 4 34 43 is) 3 1
(433)
grynea 1 29 42 2h 2;
(309 )
praeclara 2 24 59 tS
(46)
gracilis & sordida 6 29 29 Bye) Il
(166)
unijuga 8 36 18 Wy) LS 6 5
( 66 )
amica I 25 B15} Dil 10 1
(416)
palaeogama 34 25 20 18) 8 -
(906)
dejecta 35 43 20 Dy
(40)
badia On 49 itil 3
( 57)
relicta 36 AO 19 3 il
ubnata 26 32 24 18

,A
( .44

Peal

idith 6 80 14

than 10 records not included.

VoLuME 24, NuMBER 2 115

TABLE 5. Continued.

Percent Occurrence

: July August September October

Species! se ee ee Eee et Oh. = ce ABS ey eae Fa

(N) 1-15 16-31 1-15 16-31 1-15 16-30 [=i 16=3e
serena 10 36 29 24 it

(103)
residua 5 Sir 30 19 8 1
(ess)
concumbens 2, 30 36 al 9 2)
(385)
retecta 1 16 42, 24 13 3

(401)
cerogama 7 Th fol 14

(14)
parta 3 oo 21 a 9

(34)
obscura IL 8 32 39 1LY/ 3 if
(130)
cara 18 36 29 9 9
(126)
flebilis iter 58 11 al

(19)
neogama 10 34 32 iL) 6)
(339)
amatrix 26 18 50

(38)
habilis = LS 36 Al rd 1
(661)

1 Species with less than 10 records not included.

The wide annual variation in total Catocala individuals taken at any one
locality seems largely due to dramatic fluctuations in the abundance of a
very few common species (e.g. C. ilia and C. ultronia in Area 1; C. pa-
laeogama, C. residua, and C. habilis in Area 2). Weather is another factor
which undoubtedly influences records of annual abundance. It is well
known among collectors that Catocala are more frequently taken on warm,
humid nights; and thus the number of such nights during a summer will
affect the total number of Catocala recorded. Data on temperature and
humidity with respect to Catocala abundance will be obtained in our
areas in subsequent years.

Seasonal Occurrence. The seasonal appearance of adult Catocala in
Areas 1 and 2 seemed virtually identical. For example, of the 30 species
common to both areas, 17 had earliest records of capture from Washington,
and 12 had such records from Amherst (in one species the earliest re-

116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

corded date was the same in both areas). Data on the early occurrence of
the six species for which there are 50 or more records from each area are
summarized in Table 4. (Late occurrences of the species could not be as
meaningfully compared across areas because late season collecting was
much less complete in Area | than in Area 2.)

A summary of the seasonal occurrence of the Catocala species is pre-
sented in Table 5. These data were obtained by summing across years
and areas, a procedure which tends to extend the apparent season of each
species to some extent. Nonetheless, it is clear that seasonal isolation alone
is insufficient to separate most of the Catocala species from one another
(all of the species, for example, may occur during the latter half of
August). Some species may occur over the entire summer (e.g. C. unijuga,
with extreme dates of 8 July and 9 October), and these certainly must be
isolated from other species by factors other than seasonal occurrence.
However, it seems equally clear that seasonal offset may coact with other
factors in effectively isolating certain closely related pairs of species (e.g.
Dlandula and mira, serena and habilis, dejecta and retecta, subnata and
neogama).

Summary

Daily records of aduit Catocala have been kept over several years at
two localities in southern New England. For the period 1961-1969, nearly
5,000 individuals of 39 species were recorded. These data have been
summarized here in an attempt to establish the abundance, and fluctua-
tions in abundance, as well as the seasonal limits, of the species. In ad-
dition, suggestions relating to the differences in species composition at
the two localities have been advanced.

Acknowledgments

This report was written while the first author was a visiting fellow in
the laboratory of Dr. Charles L. Remington at Yale University. The
stimulation of discussions with Dr. Remington, and his suggestions regard-
ing this manuscript, are gratefully acknowledged.

some of the Catocala records from the Amherst area were contributed
by Dr. Ronald R. Keiper (1967-68) and Charles G. Kellogg (1969), and
their cooperation and assistance is greatly appreciated.

Literature Cited

Jan S | aa Me ININTOTTO? } _

3ARNES, W. AND J. McDunnovucu. 1918. Illustrations of the North American species
of the genus Catocala. Mem. Amer Mus. Nat. Hist. vol. 3, part 1

21 2 7 2 . . . a :

}ROWER, A. E. 19 30. An experiment in marking moths and finding them again
( Lepid.: Noctuidae ). Entomol. News 41: 10-15.

BUNKER, R 874. Notes ollect ’
PUNKER, Kt, 1874. Notes on collecting Catocalas. Canad. Entomol. 6: 25-26.

VoLuME 24, NUMBER 2 WIE

ForseEs, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III. Noc-
tuidae. Cornell Univ. Agri. Exp. St. Memoir 329.

FreENcuH, G. H. 1880. Notes on Catocala hunting. Canad. Entomol. 12: 241-242.

Grote, A. R. 1872. On the North American species of Catocala. Trans. Amer. Entomol.
Soc. 4: 1-20.

1873. On the genus Catocala. Canad. Entomol. 5: 161-164.

1876. On species of Catocala. Canad. Entomol. 8: 229-232.

Hampson, G. F. 1910. Catalogue of the Lepidoptera Phalaenae in the British Mu-
seum. Vol. 9. London.

HoitiaAnp, W. J. 1903. The Moth Book. New York. (Reprinted 1968, with annota-
tions by A. E. Brower).

Hutst, G. D. 1880. Remarks upon the genus Catocala with a catalogue of species and
accompanying notes. Bull. Brooklyn Entomol. Soc. 3: 2—13.

1884. The genus Catocala. Bull. Brooklyn Entomol. Soc. 7: 13-56.

Jounson, J. S. 1882. Catocalae taken in the vicinity of Frankford, Pennsylvania.
Canad. Entomol. 14: 59-60.

KETTLEWELL, H. B. D. 1958. The importance of the microenvironment to evolution-
ary trends in the Lepidoptera. Entomologist 91: 214-224.

McDunnoucu, J. 1938. Checklist of the Lepidoptera of Canada and the United
States of America. Part 1. Macrolepidoptera. Mem. Southern Calif. Acad. Sci.
Volk 1.

Row ey, R. R. aAnp L. Berry. 1909. Notes on the study of some Iowa Catocalae.
Entomol. News 20: 12-18.

SARGENT, T. D. AND R. R. Kereer. 1969. Behavioral adaptations of cryptic moths. I.
Preliminary studies on bark-like species. J. Lepid. Soc. 23: 1-9.

COMMUNAL ROOSTING IN COLIAS AND PHOEBIS (PIERIDAE)

Harry K. CLENcH

Carnegie Museum, Pittsburgh, Penna.

Two instances of communal roosting in coliadine pierids have recently
come to my attention. They have to do with different genera, of consider-
ably different body size, and occurring in different major environments,
but there are several striking similarities and I believe a common explana-
tion may satisfy both.

Colias eurytheme Boisduval

In an open meadow near Pittsburgh, Pennsylvania, at about 5:30 PM,
EST, on 18 September 1969, I saw Colias eurytheme in fairly large num-
bers preparing for the night. A few of them were still active, but most
had already sought roosts. The sun was within 15—20° of the true horizon,
but was destined to disappear a little prematurely behind a low, tree-
covered hill, and the field was already partly shaded.

118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

A few eurytheme were down in the dense grass, probably within 6-8
inches of the ground, each some distance from any other. Some, similarly
isolated, were low (less than a foot above the ground) in dense crown
vetch on a steep, still sunlit, west-facing slope that forms the eastern edge
of the field. Still others, also alone, were hanging from leaves of isolated
emergent forbs, mostly goldenrod, usually within 1-2 feet of the ground.

I saw, however, two conspicuous aggregations. Each consisted of 5
males, clinging to the leaves of a clump (2 or 3 stalks) of emergent forbs
in the middle of the field, the sun still on them. In each clump the
individuals of eurytheme were only an inch or so apart and within a foot
or two of the ground.

All the individuals I could examine closely enough, both those seen
singly and those in the two aggregations, were basking laterally (wings
appressed, the sun irradiating one side), the only basking posture I have
ever seen in Colias. The orientation to the sun was so consistent that the
best way to search for them was to keep the sun at my back as I looked:
the little patches of sunlit yellow were conspicuous, even though some
turned out on closer inspection to be merely yellowed leaves.

Phoebis sennae eubule Linnaeus

At Horseshoe Beach, Dixie Co., Florida, on 24 November 1969, at about
3:55 PM, EST, I saw an assemblage of Phoebis sennae eubule. In an
empty trailer park consisting of bare sandy ground there were a few
scattered low trees, mostly Sabal Palm. The trunk of one of these palms
was nearly covered with a dense growth of a vine part of which, about six
feet above the ground, was in the form of a hollow pocket about two feet
across, facing west. In this pocket were 11 sennae (98,22). The sun was
low and much of the area was already shaded by some nearby buildings,
although the sennae were still sunlit. Elsewhere, within a hundred feet
or so, several sennae were still actively flying along with many other
butterflies. Within the pocket of the vine, however, they were almost
all in repose, wings appressed and broadside to the sun in the same lateral
basking posture seen in Colias. There was one exception. When I ap-
proached the group one of the females had her wings fully outspread and
and her abdomen elevated at right angles to the rest of her body and to
the plane of her wings. This is known to be a rejection posture, a response
to an attending male indicating that she is not receptive to copulation.
The rest of the group was quiet, however, and she soon abandoned the

nd folded her wings together like the others.
‘en or fifteen minutes earlier I had looked briefly at the as-
had noted only four or five individuals in it. I therefore

VoLUME 24, NuMBER 2 119

suspect that the group was slowly being added to by new arrivals. ( Pos-
sibly one of these was a male who on arriving had attempted to copulate
with the female and had elicited the rejection posture I saw.)

The color of sennae eubule is distinctly greenish yellow. The resem-
blance of the color of these individuals to the yellow-green leaves of the
vine was so extremely exact that it was difficult to distinguish one from the
other, even at close range.

Discussion

Nearly all the roosting Colias I saw, whether they were single or in the
aggregations, were situated more or less close to yellowed or to yellowing
leaves, whose color closely matched their own. An association of Colias
with yellow flowers or leaves has long been known (Scudder 1889, p.
1123; Clark 1932, p. 159).

Several times I have noticed a similar association in Phoebis. In a local-
ity in eastern Mexico I once saw them at rest in a low shrub with many
yellow leaves. The size and color of these leaves so closely resembled the
Phoebis that at a short distance it was impossible to tell which was which.
At Chokoloskee, Collier Co., Florida, on 20 November 1969, I watched a
Phoebis agarithe maxima Neumoegen in late morning, basking laterally
for several minutes on a yellowed leaf of papaya along a path in a patch of
low forest. This leaf was many times the size of the agarithe, but its
orange yellow color was so close a match that the butterfly was almost
invisible against it. This latter observation is particularly suggestive when
taken in conjunction with the sennae already described. These two species
are of about the same size but they are conspicuously differently colored:
greenish-yellow in sennae, orange-yellow in agarithe. In each instance the
background color closely resembled that of the particular species. See
also Longstaff (1909, pp. 640-643).

Communal roosting is not well-known in either Colias or Phoebis. I
think that in both genera it is an occasional occurrence only. This assump-
tion is supported in Colias by the fact that at the same time that I saw the
aggregations there were many other individuals preparing to roost alone.
The above observations suggest the possibility that among the various
factors that enter into the choice of a roosting site, the presence nearby
of an appropriate shade of yellow is important. This may lead to single
roosting individuals, but on the other hand, if the individual itself im-
proves the color match or adds to the conspicuousness of the yellow at
the site, then it may attract still others and result in an aggregation. In
this sense, then, communal roosting may be simply a by-product of their
xanthotropism.

120 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Literature Cited

Crark, Austin H. 1932. The butterflies of the District of Columbia and vicinity. U.
S. Nat. Mus. Bull. 157. 337 pp.

Loncstarr, G. B. 1909. Bionomic notes on butterflies. Trans. Ent. Soc. London
(1908). pp. 607-673.

ScuppEr, SAMUEL H. 1889. The butterflies of the eastern United States with special
reference to New England. Cambridge, Mass. 3 vols.

TWO NEW MEXICAN HESPERIIDAE

Lez D. MILLER
The Allyn Foundation, Suite 712, Sarasota Bank Building, Sarasota, Florida

Among the Hesperiidae collected by the Carnegie Museum-Catholic University of
America expedition to eastern Mexico in January, 1966, were two short series of ap-
parently new hesperiine skippers. When the Allyn Mexican expedition was under-
taken in January and February, 1969, an opportunity was provided to obtain more
specimens and to observe more closely the habits of these hesperiids. The combined
series from the two trips comprise the type-series for the two new species below.

Although perhaps not so many specimens nor species can be taken in Mexico during
the temperate zone winter as might be collected during the summer months, many
worthwhile species apparently are found only during January and February, or at

least are most abundant during this time. These two new skippers are evidently among
such “winter” butterflies.

Vinius freemani Miller, new species
Higs, 2703), 312) (Gucenutalnap)

Male: Head, thorax and abdomen blackish-brown, clothed above with greenish-
ochre hairs; below with dense tan hairs. Antennae brown above, ringed with brown
and bright fulvous below; nudum dull brown. Palpi brownish-fulvous above, grayish-
tan below. Legs densely clothed with brownish-fulvous hairs.

Upper surface of forewing reddish-fulvous with veins fuscous and with the follow-
ing fuscous markings: apex and outer margin broadly dark; this border narrowing in
space M.-M., then broadening to anal angle; the border enclosing three light subapical
spots in spaces R—-Ri, Ru-R; and R;-M:. A dark streak bisecting discal cell through-
out its length. A patch just distad of the cell in Mi—M> connecting the outer marginal
band with the black androconial mass; latter passing along bases of spaces M--Cu: and
Cuz, then diagonally through space Cu:-2A to 2A.

Cu
‘lind wing fuscous above, densely overlaid with basal brownish hairs extending
he anal veins; a diffuse reddish-fulvous cell-spot; a short extradiscal band of
Pare. i small spot in space Rs—Mh, a long one in M:—M; and some-
an : M:-Cu, and Cu:-Cue. Fringes fulvous, darker from
10 AEX 01 d plain or only slightly checkered.
broadly black basally and along anal cell to inner angle,
s a tapering submarginal band, broad in space Cus—
ah te M:-M: and M:—Mz cells. Remainder of wing dull
upper surface lightly indicated by darker fulvous.

alone '

VoLUME 24, NuMBER 2 1A

Figs. 1-6. New Hesperiidae from Mexico. 1, Vinius freemani Miller, holotype ¢,
upper surface. 2, Same, under surface. 3, Vinius freemani Miller, paratype 9, VERA-
CRUZ: 2 mi. SE Coatzacoalcos, upper surface. 4, Virga clenchi Miller, holotype 4,
upper surface. 5, Same, under surface. 6, Virga clenchi Miller, paratype 9, TA-
BASCO: 2—3 mi. E LaVenta, upper surface. The figures are approximately twice life-
size.

Under side hind wing fulvous, shaded with fuscous in space 2A—3A, and with an
irregular fuscous spotband across discal cell from Sc—Rs to vein 2A (the spot in Sc—
Rs distad of the others), and a V-shaped band of small fuscous spots in the same
spaces, the one in M:—Ms nearest outer margin. Fringes fulvous, slightly darker
toward apex of forewing, and slightly checkered with fuscous.

Forewing length of holotype, 12.0 mm., of the seven male paratypes, 11.5 to 12.5
mm., averaging 11.9 mm.

Male genitalia (Fig. 7) quite distinct from those of other Vinius species (see Hay-
ward, 1950, pl. 10, fig. 13, and Evans, 1955, pl. 58), with simpler valvae and tip of
penis with a long saw-like ventral projection.

Female: Body and appendages as in male, except that dorsal hairs darker, and
ventral ones somewhat duller tan.

Upper side of forewing fuscous with dull fulvous markings as follows: two small
spots near end of discal cell, the lower one somewhat the larger; three subapical spots
as in male; an extradiscal band of spots beginning as a small dot in space M:i—M> mar-
ginad of subapical spots and proceeding posteriad and somewhat basad to a V-shaped
spot in Cus-2A. Hind wing likewise fuscous above with a poorly defined dull fulvous
cell-spot and a row of extradiscal fulvous spots as in male, but composed of smaller
spots. Fringes light brown at anal angle of hind wing, shading to dark brown at
forewing apex. Under surface like that of male, but darker and duller.

Ten female paratypes with forewing lengths of 11.5 to 13.5 mm., averaging 12.6
mm.

Described from eighteen specimens, eight males and ten females, from two localities
near Coatzacoalcos, Veracruz, Mexico.

HOLOTYPE ¢: MEXICO: VERACRUZ: 2 mi. SE Coatzacoalcos, Sta. 24, grassy
scrub/marsh/palmetto savanna association; 18-i-1966; H. K. Clench and L. D. Miller

129 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(L. D. Miller specimen no. 1966-738 ); CM-CUA expedition.

PARATYPES: 62 79, same locality and date as holotype. 16 39, MEXICO:
VERACRUZ: 1-2 mi. W Nanchital, sea level, savanna; 2-ii-1969, Sta. 19 (1¢ 29),
3-ii-1969, Sta. 21 (19); L. D. and J. Y. Miller; Allyn Mexican expedition.

Holotype and 4é and 6@ paratypes in Carnegie Museum, 28 and 29 paratypes
in Allyn collection, 1g and 19 paratypes in collection of H. A. Freeman and 19
paratype in collection of the Direccion General de la Faune Silvestre, Mexico, D. F.

I take great pleasure in naming this distinctive skipper for my good
friend H. A. Freeman of Garland, Texas, in recognition of his magnificent
work on the Hesperiidae of Mexico.

V. freemani will key to letis (Plétz) in Evans (1955, pp. 71-72). The
under side of the hind wing of female freemani, however, is not noticeably
greenish as in letis. Since letis is known only from southeastern Brazil,
Paraguay and Argentina its conspecificity with the Mexican insect would
be suspect, but the differences between the genitalia of freemani and letis
are great, much greater than one could expect from the superficial
similarities. V. sagitta (Mabille) from the Chiriqui, Panama, is clear yel-
low below, though marked rather like freemani, and the genitalia are dif-
ferent, whereas V. t. tryhana Kaye, which is known from Mexico, is very
different superficially and genitalically.

I suspect that freemani has been overlooked by collectors since it looks
and behaves rather like a small reddish version of the abundant Hylephila
phyleus phyleus (Drury). Both species show a distinct preference for
various white flowers in the Coatzacoalcos area and may be seen in
swarms on suitable blossoms, occasionally a hundred or more phyleus and
one or two freemani on a single bush. My wife and I collected more or less
extensively in a number of other areas around Coatzacoalcos without find-

ing freemani, and I believe this skipper is restricted to swampy savanna
situations.

Virga clenchi Miller, new species
Figs.4,5(3 ),6(92),8(64 genitalia)

Male: Wead, thorax and abdomen blackish-brown clothed above with dark-brown

hairs, below with tan hairs. Antennal shaft brown above, yellow and brown ringed
below; club brown above and yellow below; nudum dull brown. Palpi brown above,
clothed with tan hairs below. Legs clothed with tan hairs.

Upper surface of forewing fuscous with a bronze cast and with fairly prominent
bronze-fulvous extradiscal spots near bases of cells M.—Cu, and Cui-Cuz,_ a bronze-
fulvous streak in space Cu.x-2A and some fulvous costal shading. In some specimens
two or three light subapical spots vaguely indicated.

Hind wing above fuscous with a bronze sheen and an ill-defined band of ex-
tradiscal streaks from cell M:—M. to cell Cu,—2A. Fringes gray-brown with fuscous
checkering on forewing.

Under surface of forewing fuscous with markings in M;-Cui and Cu;-Cuz repeated

in grayis oe ig Se to Cu, yellowish distally and small gray subapical spots in
spaces Ki—M, and M.—M..

VoLUME 24, NuMBER 2 123

1 mm

Figs. 7-8. ¢@ genitalia of new Mexican Hesperiidae. 7, Vinius freemani Miller,
genitalia of paratype 6, VERACRUZ: 2 mi. SE Coatzacoalcos (LDM slide M-1577).
8, Virga clenchi Miller, genitalia of paratype 6, VERACRUZ: 2 mi. SE Coatzacoal-
cos (LDM slide M-1581).

Hind wing below fuscous with basal grayish-tan area and an extradiscal band of
same color from costa to Cu,-—2A; veins yellowish-gray. Fringes grayish-tan, those of
forewing checkered with fuscous.

Forewing length of holotype, 9.0 mm., of the ten male paratypes, 8.5 to 9.5 mm.,
averaging 9.0 mm.

Male genitalia (Fig. 8) quite different from those of other two species known from
Mexico and Central America, virginius (Moschler) and xantho (Schaus), figured by
Evans (1955, pl. 59) and Godman and Salvin (1879-1901, pl. 103). V. clenchi
especially characterized by simple valvae and long gnathos.

Female: Body and appendages as in male.

Upper surface of forewing fuscous with no bronze sheen and with bronze-fulvous
spots in four of five specimens in spaces M;—Cu: and Cu;—Cuz and two subapical
bronze spots in two of five specimens.

Hind wing above fuscous with poorly defined extradiscal bronze band as in male.
Fringes as in male, but checkering more pronounced.

Under surface as in male.

Five female paratypes with forewing lengths of 9.0 to 10.0 mm., averaging 9.6
mm.

Described from sixteen specimens, eleven males and five females, from various |o-
calities in coastal southern Veracruz and northernmost Tabasco, Mexico.

HOLOTYPE ¢: MEXICO: VERACRUZ: 2 mi. SE Coatzacoalcos, Sta. 24, grassy
scrub/marsh/palmetto savanna association; 18-i-1966; H. K. Clench and L. D. Miller
(L. D. Miller specimen no. 1966-717 ); CM-CUA expedition.

PARATYPES: MEXICO: VERACRUZ: 26, 13 mi. NW Alvarado, 10 m.; Sta.
23 (CM-CUA); 16-i-1966; H. K. Clench and L. D. Miller (1¢); Sta. 12 (Allyn
Mexican); 29-i-1969; L. D. and J. Y. Miller (1¢). 19, 16 mi. E Acayucan, Sta. 25
(CM-CUA); 18-i-1966; H. K. Clench and L. D. Miller. 26 39, same locality and
date as holotype. 1¢, 1-2 mi. W Nanchital, sea level savanna, Sta. 17 (Allyn Mexi-
can); 1-ii-1969; L. D. and J. Y. Miller. 16, 5 mi. SE Coatzacoalcos, 10 m., dense
scrubby swampland, Sta. 20 (Allyn Mexican); 3-ii-1969; L. D. and J. Y. Mill

124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABASCO: 42 19, 2-3 mi. E LaVenta, 10 m., swamp forest, Sta. 15 (Allyn Mexi-
can); 31-i-1969; L. D. and J. Y. Miller.

Holotype and 3é and 3@ paratypes in Carnegie Museum, 5¢ and 19 paratypes
in Allyn collection, 1¢ and 1@ paratypes in collection of H. A. Freeman and 1 ¢
paratype in collection of the Direccion General de la Faune Silvestre, Mexico, D. F.

I am happy to name this little skipper for my friend and colleague
Harry K. Clench of Carnegie Museum, Pittsburgh, Pennsylvania, who
helped collect those specimens of the type-series collected by the
Carnegie Museum—Catholic University of America expedition in 1966.

V. clenchi may have masqueraded in collections as virginius from which
it may be distinguished by its smaller size, less prominent markings and
the genitalic differences (for comparison see particularly the figure of the
genitalia of virginius, as cometho Godman, in Godman and Salvin, 1879-
1901, pl. 103). The present species is found in small numbers in a rather
broad area along the southern coast of the Gulf of Mexico, usually in
moist or swampy habitats, but its occurrence in a disturbed situation east
of Acayucan demonstrates that clenchi is an adaptable insect. This Acayu-
can locality is not, however, far from the great marsh west of Coatzacoal-
cos, so the specimen might have strayed from a more suitable habitat.
These butterflies are avid flower visitors, preferring white blooms, as do
most skippers.

Finally, I would like to thank H. A. Freeman for checking these two
skippers against his great Mexican hesperiid collection.

References Cited

Evans, W. H. 1955. A catalogue of the American Hesperiidae. Part IV. London, 499
pp.

GopMan, F.. D., anp O. Savin. 1879-1901. Biologia Centrali-Americana. Insecta.
Lepidoptera-Rhopalocera. London, 2 vols.

Haywarp, K. J. 1950. Genera et species animalium Argentinorum. Tomus secundus.
Insecta, Lepidoptera (Rhopalocera), familia Hesperiidarum, subfamilia Hesperi-
inarum. Buenos Aires, 388 pp.

VoLUME 24, NuMBER 2 125

VARIATIONS IN THE MARKINGS OF PIERIS RAPAE (PIERIDAE )
INDUCED DURING THE PUPAL STAGE

JoHNn M. KoLyeEr
55 Chimney Ridge Drive, Convent, New Jersey, U.S.A.

Introduction

The present work represents the conclusion of a study which has
involved observations on the effects of feeding larvae certain chemicals
(Kolyer, 1966) and on the effects of environmental factors (Kolyer, 1969)
on the wing markings of the European cabbage butterfly Pieris rapae
(Linnaeus). Specific objects were to note variations in markings as a
result of (1) reduced-temperature storage of pupae of various ages, as an
extension of previous work (Kolyer, 1969), (2) brief, barely sublethal
heating of pupae, and (3) exposure of pupae to short-wave ultraviolet
(UV) radiation. Incidental observations, e.g. on larval disease and the
yellow form of P. rapae, also are included.

Experimental Procedures

Rearing. Final instar larvae were reared on an artificial diet at 79 +
2°F and 45 + 5% relative humidity under continuous cool white fluores-
cent light. Larval development was completed on cabbage leaves from
refrigerated heads (see Kolyer, 1966); rearing room conditions were: 70-
81°F, approx. 20-33% rel. hum., diffuse sunlight (March 6—April 17, 1969,
Convent, New Jersey ).

Refrigeration of pupae. As in previous work (Kolyer, 1969), pupae
were packed carefully along with facial tissues in fiber mailing cans, which
were covered with polyethylene bags to exclude moisture and stored in a
refrigerator at 32-36°F for 154-156 days (approx. 5 months).

Heat treatment of pupae. Two test tubes, one inside the other, were
immersed in hot water in a Dewar (vacuum-jacketed ) vessel and allowed
to equilibrate so that the temperature of the air within the inner tube ap-
proximated the temperature of the bath. Then pupae were added to the
inner tube and allowed to remain for the specified time periods.

UV irradiation of pupae. Pupae were exposed, righthand wing case up,
at a distance of 2 inches from an 8-watt glow discharge (mercury) lamp
bulb giving light principally at 254 millimicrons. The intensity is given as
17 microwatts/cm.? at 1 meter, which is “sufficient for effective air, sur-
face, and liquid disinfection” (Anonymous, 1965). Temperature rise of a
mercury thermometer bulb 2 inches from the lamp was considered
negligible (0.5° C after 40 min. exposure ).

126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Results and Discussion

Variability of markings. The great variation in markings among individ-
uals of P. rapae was mentioned in the earlier work. Wing length of this
species varies from year to year (Petersen, 1947), and such variation
also may be true of the proportion of “immaculata” (spotless form) males
in the spring brood. Of 21 males taken at Flemington, New Jersey, on May
1, 1965, 11 were “immaculata” by the arbitrary criterion of less than 10
black scales in the forewing spot (Kolyer, 1966). In contrast, only 4 of
28 males taken on April 21-27, 1969, at Morristown, New Jersey (29 miles
northeast of Flemington) were “immaculata” (No. 1, Plate I). Refrigera-
tion of nondiapause pupae at 34-39°F for 5 months gave 5 “immaculata” of
21 males (Kolyer, 1969), thus surpassing natural conditions in producing
this form in at least one instance.

Controls—deformed head, yellow form. The control pupae were allowed
to remain under rearing room conditions (cited above), and all adults
eclosed in 9-12 days. Adult forewings are shown in No. 3, Plate I, and
Nos. 5 and 6, Plate II.

The first control group (No. 3, Plate 1) included an otherwise normal
female with undeveloped head (no proboscis, palpi, or antennae), shown
in No. 4, Plate I. A similar result was noted with a male adult fed the dye
safranin bluish as a larva (Kolyer, 1965).

During completion of rearing of the second control group (Nos. 5 and
6, Plate II), a yellow-green larva, markedly different in shade from the
other (grass-green) larvae, was isolated and reared to pupation. In 9
days a yellow-form male eclosed (the complete specimen in No. 5, Plate
II). Since this was the only yellow form among 149 adults which even-
tually eclosed from the batch of larvae in question, the probability of hav-
ing picked the yellow-form larva by chance alone was only 1.3%. A
recessive yellow form of the larva of Pieris napi (L.) has been described
(Gladman, 1962), but adults in this case are normal in appearance.
Incidentally, the complete specimen ( male) in No. 3, Plate I, also is a
yellow form.

Larval disease. A minority of larvae evidenced black spots (appearing
under magnification as black-rimmed pits) on the integument. Two of
these larvae were sent to Mr. G. M. Thomas of the University of California
‘see Acknowledgments) for a disease diagnosis. The larvae were found

uffer a bacteriosis caused by a strain of the rather common insect
sen Serratia marcescens (Bizio). This strain was nonchromogenic
ind association of the disease with the black spots is specula-

ots on larvae usually are symptomatic of microsporidian

VoLUME 24, NuMBER 2 DAF!

>

PLATE I

Specimens of Pieris rapae (L.). 1, 2, Forewings (and underside of hind wings in
bottom rows) of series taken at Morristown, New Jersey, on April 21-27, 1969; 1,
$6; 2, 2 2; 3, control group (pupae developed at 70-81° F') for Nos. 7-9, Plate II;
4, 2 from preceding control group with undeveloped head, photographed through
16X microscope.

128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

infections, but no protozoans were found, nor was there evidence of virus,
fungi, or nematodes.

Some of the spotted larvae died, or yielded deformed pupae which
died, but others gave normal pupae and adults. One adult was sent to
Mr. Thomas. Thorough examination of the tissues and blood revealed no
microbial etiology; a complete analysis showed no bacteria, protozoans,
virus, fungi, or nematodes.

Refrigeration of pupae. In the previous work (Kolyer, 1969), non-
diapause pupae were refrigerated (34-39°F) at an age of 12-24 hours
from pupation (final molt); after 5 months these pupae were allowed to
develop at room temperature and yielded adults with significantly reduced
(spring brood ) markings.

In the present study, pupae of various ages were refrigerated (32-36°F )
for 5 months with the objective of gaining information on the time of
determination of the markings. In the meal moth Ephestia kitihniella
(Zeller), and in Lepidoptera in general, the wing pattern is completely
determined early in the pupal stage (Magnussen, 1933, and Pohley, 1959).
In P. rapae pupae the spots have been determined at least by the time
of visible deposition of white pigments (pupal age about 135 hours), as
shown by the lack of white pigment in the scales within the spot at this
time.

Refrigeration results are given in Table 1; two control groups are
involved because the larvae had been received from the Department of
Agriculture in two separate batches about one month apart. From the
24-34 hour old pupae, eight of twelve normally-expanded males were
“immaculata.” Refrigerated females tended to lack the apical marking
and to have weak spots; in one case both spots and apical marking were
almost completely absent (middle wing in 4th row, No. 10, Plate III).
Melanization of the underside of the hind wings, as in the spring brood,
also occurred (see No. 10, Plate III).

The conclusion is that these nondiapause pupae had to be refrigerated
at an age of less than 48 hours to allow suspension of development and
eventual eclosion. Also, the lack of reduction of markings on the pupal
wings of the 48-56 hour old group (No. 9, Plate IL) suggests that the
markings have been determined by 48 hours. Concurrent studies indicate
that the scales develop at some point between 27 and 87 hours, and it may
be th. it the fate of a scale to be white or black must be decided before the

’s out from the original stem cell. (As described by Lipp (1957),
Le of the freshly-molted P. brassicae pupa is composed

i of stem cells which later give the scale and socket arrange-
‘expectedly, one male in the 133-143 hour old group managed

VoLUME 24, NuMBER 2 129

Provera) UT

Specimens of Pieris rapae (L.). 5, 6, Forewings (and underside of hind wings at
bottom) of control group (pupae developed at 70-81° F) for Nos. 10-15, Plate III;
5, 64; 6, 2 9; 7-9, forewings of adults from pupae refrigerated at 32-36° F for 5
months, pupae refrigerated at specified time from pupation: 7, 10-18 hours; 8, 24—34
hours; 9, 48-56 hours.

130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Data on pupae refrigerated for five months.

ee Age ERS, No. Eclosed*
ontro
oe Group No. Fully Expanded Crum- No.
frigerated in Plates Males Females pled Died* Notes

10-18 3 FU) 4 3 17 Many dead pupae showed
light markings.

24-34 3 3 (2) 2 4 PAN Ditto.

24—34 5&6 SE) 7 19 Ditto.

48-56 3 0 0 0 29 Some pupal wings showed
markings—essentially
like summer brood.

78-88 3 0 0 0 29 —

80-86 5&6 0 0 0 bil Most had black wing cases.

103-109 5&6 0 0 0 pill Some with black wing

cases; 5 showed markings
—like summer brood.

120-130 8) 0 0 0 ult a

133-1434 5&6 LO) © 0 48 One with black wing cases;
8 showed markings—
like summer brood.

Controls 3,5&6 total of 72 2 i —

1 Eclosion was completed within 11 days after removal to rearing room conditions 71—83° F, 50—
57% rel. hum. ).

2 The number of “immaculata’”’ form is given in parentheses.

% Many died after the pupal wing markings had developed. In the case of several pupae (total of
11 for entire column) the adult split the pupal case but failed to emerge.

4 The wing cases were white (pigment deposited ).

to eclose (No. 11, Plate III); the markings seemed not significantly less
intense than those of the control males.

Heat treatment of pupae. A fatal high temperature for butterfly pupae
has been said to be 39-42° C (Uvarov, 1931), but Kiihn (1936) subjected
meal moth pupae of various ages to 45° C for 45 minutes to observe effect
on the markings, and Schrader (1929) exposed pupae of Vanessa carye
( Hiibner ) to 48° C for “a short time” to cause aberrations.

In the present work, pupae of various ages (from 10 to 128 hours) were
exposed to air under various conditions within the limits of 36-48° C and
10-30 minutes. Conditions under which all pupae died were (number of
pupae, pupal age in hours, temp. in ° C, time in min.): 6, 10-18, 45.3-48.3,
20; 2, 21-27, 43.8-47.0, 20; 4, 23-39, 43.3-46.6, 30. Among conditions which
allowed all pupae to live and yield adults were: 4, 13-18, 36.3-39.1, 30; 7,
19-25, 38.9-45.0, 10; 4, 22-28, 41.6-46.4, 20; 2, 24-39, 41.8-45.3, 25. These

aS

ns of Pieris rapae (L. ). 10, Forewings (and undersides of hind wings for 2
0) of 6 4 (first 3 rows) and 9 9 from pupae refrigerated (at 24-34
upation) at 32-36° F for 5 months; 11, forewing of ¢ from pupa

VoLUME 24, NuMBER 2 (i

PLATE III

similarly refrigerated at 133-143 hours from pupation; 12, 13, 2 2 from pupae which
had right wing cases exposed to UV light (see text); 12, 12 hours exposure; 13, 36
hours exposure; 14, 15, wings of adults from pupae exposed to air at 41.4—47.5° C for
20 min. (see text); 14, largely-scaleless forewing of 2, photographed through a 16x
microscope against a black background; 15, forewing of ¢ with side-lighting to illus-
trate “bumpy” surface, photographed through 16 microscope.

132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

results suggest that surviving pupae had been exposed to very nearly
lethal conditions. However, normal specimens with summer brood mark-
ings (like the controls ) resulted in almost all instances.

Pronounced structural abnormalities occurred in only one case (con-
ditions: 14, 74-84, 41.4-47.5, 20). Two pupae died, one adult split the
pupal case but failed to eclose, 3 pulled partly out of the case, 7 had
crumpled wings, and only one (male) expanded normally. One female had
particularly wrinkled and distorted wings with a high degree of scaleless-
ness, but the markings (spots) still were visible; No. 14, Plate III, shows
the forewing in the region of the spot photographed against a black back-
ground. The normally-expanded male had “rough” wings on close ex-
amination, due to erection of isolated patches of scales. The other in-
dividuals had more or less crumpled wings, in some cases with an unusual
“bumpy appearance though well clothed with scales (as illustrated in No.
15, Plate IIT).

Dehydration has been suggested to explain changes in meal moth wing
pattern caused by heating pupae at about 45° C for 45 minutes (Braun,
1939), but the weight loss for the P. rapae pupae heated at 41.4—47.5° C for
20 min. was only 0.3% (14 pupae weighed 2.185 grams before heating,
2.178 grams after heating ).

Scale loss as a result of heating pupae has been noted for the meal moth
(Kuhn and Merkel, 1955). Also, the present results, especially for the
female shown in No. 14, Plate III, are reminiscent of the effects, including
wrinkling and scalelessness, reported for Papilio pupae subjected to beta
rays (Kishi, Miwa, and Mori, 1942). In conclusion, heating P. rapae
pupae failed to change the distribution of melanin but did, in one case,
cause considerable structural deformity.

UV irradiation of pupae. Young pupae of the meal moth were irradiated
with UV light from a mercury lamp by Kohler (1941) to give disturbances
in scale arrangement and transformations in pattern.

Thirty-six P. rapae pupae, 50-60 hours old, were placed under the UV
lamp as described under “Experimental Procedures” above. Twelve were
removed from the light after 2 hours exposure, twelve more after 12 hours
total exposure, and the rest after 36 hours total exposure. Results were

(exposure time in hours, males expanded normally, females expanded
normally, individuals with right wing crumpled, individuals with both
wings crumpled ): 2, 4, 2, 7, 1: L2, 3, 3; 6, O36" a ormle (Of 45 controls,
all eclosed, and only 2 had crumpled wings.) The adults eclosed 8-10 days
from pupation (conditions: 70-8]° F’, 28-33% rel. hum.), with the excep-

M ‘ 7 P| 3) eC 1
tion of one female which eclosed about 25 days from pupation.

Three le rmallveeyn: : *
Three of the normally-expanded females had notably asymmetric

VoLUME 24, NuMBER 2 133

markings. Two (Nos. 12 and 13) are shown in Plate III. Examination of
the wings under a 16X microscope showed that the scales on the upper
surface of the forewing had a shriveled, narrowed appearance. Within
the spot the scales were essentially white, the gray appearance of the spot
being due to the presence of normally-pigmented black scales within the
spot on the lower wing surface, beneath the transparent membrane. In
these cases the effect of UV irradiation seemed to be to deform the scales
and prevent those scales in the spot from undergoing black pigmentation
(melanin formation ), a process that begins approximately 25 hours before
eclosion.

Summary

The European cabbage butterfly Pieris rapae (Linnaeus) has variable
black markings which may be much reduced in the spring brood, even to
the extent of disappearance of the forewing spot in the male (“im-
maculata” form). Attempts were made to influence these markings b’/
various treatments of nondiapause pupae originating from larvae reared at
79°F under continuous light.

Refrigeration of 24-34 hour old pupae at 32—36° F for 5 months gave a
high proportion (8 of 12) of males of the “immaculata” form and females
with much reduced apical markings and weak spots (in one case virtually
no markings at all). Reduction in markings also was achieved with 10-18
hour old pupae. However, pupae varying in age from 48 to 143 hours
(white pigment appears at about 135 hours) failed to eclose with only one
exception (a 133-143 hour old pupa, which yielded a male with un-
reduced markings), and the markings sometimes visible on the pupal
wings were little reduced. This suggests that to affect markings refrigera-
tion must precede outgrowth of the scales from the stem cells of the wing
epithelium.

Heating of pupae under barely sublethal conditions failed to cause
redistribution of melanin but produced structural deformities in one ex-
periment (74-84 hour old pupae exposed to air at 41-48° C for 20 min-
utes ); the wings were partly scaleless and wrinkled in several cases and
were highly deformed and largely scaleless in one individual.

Exposure of 50-60 hour old pupae to UV light (largely 254 milli-
microns) caused deformity of the scales in some individuals, with lack of
black pigmentation of scales within the forewing spots on the upper, but
not the under, surface of the exposed wing.

All these results seem consistent with final determination of the wing
pattern early in the pupal stage, as reported for the meal moth Ephestia
kiihniella (Zeller ) in intensive studies by Kihn and others.

134 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Acknowledgments

The author gratefully acknowledges the contribution of larvae for this
and other work by the Columbia, Missouri station of the United States
Department of Agriculture, where Mr. Benjamin Puttler was Assistant
Director and Mr. Richard K. Morrison was in charge of the insectary rear-
ing program. The author also is indebted to Mr. Gerard M. Thomas of the
Agricultural Experiment Station, College of Agricultural Sciences, Univer-
sity of California, Berkeley, for the disease diagnoses.

Literature Cited

AnonyMous, 1965. Scientific Apparatus and Reagents, Arthur H. Thomas Co.
(Catalog). A. H. Thomas Co., Philadelphia, Penna. (see Cat. No. 6322-C, p. 665).

Braun, W., 1939. Disturbances in the process of cell-division in the pupal wing of
the flour-moth Ephestia kiihniella as result of heat treatment. Cytologia 10(1/2):
40-43.

GLADMAN, J. C., 1962. Some notes on the early stages of Pieris napi (L.) including
a yellow form of the larva. Entomologist 95(1193): 253-254.

Kisut, S., M. Mrwa, & K. Mort, 1942. Influence of beta rays on the pupal wing of
butterfly. Zool. Mag. (Tokyo) 54(12): 502-505.

Kouuer, W., 1941. Experimentelle Unterssuchungen tiber die Determination des
Zeichnungsmusters bei der Mehlmotte Ephestia kiihniella Zeller. Vierteljahres-
schr. naturforsch. Ges. Zurich 86: 77-151.

Koryer, J. M., 1965. The feeding of coloring matters to Pieris rapae larvae. J.
Res. Lepidoptera 4(3): 159-172.

1966. The effect of certain environmental factors and chemicals on the markings
of Pieris rapae (Pieridae). J. Lepid. Soc. 20(1): 13-27.
1969. Effects of environmental factors on the markings of Pieris rapae (Pieridae).
J. Lepid. Soc. 23(2): 77-94.
ae A., 1936. Versuche tber die Wirkungsweise der Erbanlagen. Naturwiss. 24
1): 1-10.

Kunn, A., & A. MerKeL, 1955. Concerning some of the mutations and phenocopies
influencing the scaly covering of moth wings. Biol. Zentralbl. 74(3/4): 113-145.

lupe, C., 1957, The significance of differential cell division in the origin of the
scale pattern on the wing of Pieris brassicae. Biol. Zentralbl. 76(6): 681-700.

MAGNussEN, K., 1933, Untersuchungen zur Entwicklungsphysiologie des Schmetter-
lingsfliigels. Zeitschr. Wiss. Biol. Abt. D, Roux’ Arch. Entwicklungsmech. Organ.
128(3): 447-479.

PreTERSEN, B., 1947, Die geographische variation einiger Fenno-Skandlischer Lepidop-
teren. Zool. Bidrag Uppsala 26: 329-531.

vontby, Hi. J., 1959. On meal moth wing growth under normal and experimental

Biol. Zentralbl. 78(2): 231-250.
'929. ‘The causes of change in color patterns of butterfly aberrations.
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Insects and climate. Trans. Ent. Soc. of London 79(1): 1—

SCHRADER
,

UVvARo

VoLUME 24, NuMBER 2 135

THE BIOLOGY AND LABORATORY CULTURE OF
CHLOSYNE LACINIA GEYER (NYMPHALIDAE)

Boyce A. DruMMonp III, Guy L. BusuH, Anp THoMaAs C. EMMEL,!

Department of Zoology, University of Texas, Austin

The nymphalid butterfly, Chlosyne lacinia Geyer, is the most widely
distributed species of its genus, ranging from Argentina northward into
Texas, New Mexico, Arizona, and the Imperial Valley and adjacent desert
areas of California (Comstock, 1927; Ehrlich and Ehrlich, 1961). Oc-
casionally it may penetrate as far north as Kansas and Nebraska (Klots,
1951).

The objective of this paper is to present an outline of the life history and
laboratory rearing techniques established as a result of an investigation
recently initiated on the ecological genetics of the various color morphs
present in the Texas populations. Details concerning the inheritance and
fitness in natural populations of certain color morphs will be published
later.

Rearing

Larvae of Chlosyne lacinia were reared in the laboratory from egg
masses collected in the field or laid in the laboratory by wild-caught
females. Individual 4 and 6 quart polyethylene tubs with tight-sealing
lids were used to hold the larvae of separate broods in early rearing
procedures. Both ventilated and unventilated lids were tried. The larvae
were supplied daily with fresh leaves of Helianthus annuus and the con-
tainers cleaned of frass.

No attempt was made to regulate humidity, but larvae in unventilated
tubs in which the humidity was high (90% or more) developed more
rapidly than those in the ventilated tubs. However, at high humidity the
larvae were much more susceptible to disease; mortality was particularly
high in the fourth and fifth instars. Most containers were kept at a con-
stant 21° C under artificial light of 16 hour daylength. It was noted during
the early attempts at rearing that larvae developed more rapidly at higher
temperatures but were more subject to disease. Although the use of ven-
tilated lids reduced mortality of larvae reared at high temperature, the
Helianthus leaves dried out within a matter of hours, rendering them
inedible to the larvae and thus lengthening developmental time.

The feeding and disease problem was satisfactorily solved by employing
the Special Vanderzant-Adkisson Modified Wheat Germ Diet ( Adkisson et
al., 1960) as an artificial medium (available commercially from Nutritional

1 Present address: Department of Zoology, University of Florida, Gainesville 32601.

136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Biochemical Company, Cleveland, Ohio 44128). The medium was sup-
plemented with the antibiotics aureomycin and acromycin, as well as
methyl parasept; these were added at the concentration of 0.001%
by weight to help check disease. Finely ground dried Helianthus leaves
(2.5 to 10% by weight) were normally included as a feeding stimulant.
Although larvae have been raised on media lacking the Helianthus addi-
tives, they developed more rapidly and only acquired full coloration on the
diet supplemented with leaves, particularly at the higher concentrations.

Larvae raised on the artificial medium were normally kept at 24°—
26° C under a 16 hour daylength or under continuous lighting; the latter
condition caused some larvae to enter diapause. At this temperature the
medium, which was placed in strips on the bottom of unventilated poly-
ethylene containers, required changing only once a week for the first three
instars, and only slightly more often as the larvae increased in size.

Mortality of medium-raised larvae was low through the first four instars,
but the fifth instar larvae were quite susceptible to infection. The overall
incidence of infection, however, was much lower than in larvae reared on
leaves alone. The medium apparently fulfilled most of the nutritional re-
quirements of this butterfly since it supported three generations of insects
in our laboratories. The effects of the diet on fecundity and viability were
not established but are now under study.

Adults were collected by placing pupae in one-pint paper ice cream
cartons, the solid tops of which were replaced with gauze. Upon emer-
gence, the adults were maintained under an artificial 16 hour day and fed
once or twice a day on a solution of one-third honey (or sugar) and water
(1:3 by volume) soaked into pieces of sponge or commercially available
pressed cotton (Coets, Quilted Squares; Personal Products Company, Mill-
town, New Jersey ) placed on the gauze.

Mating was accomplished either in small wire cages under the partial
shade of a tree or in large indoor mating cages artificially lighted with a
bank of four Gro-Lux fluorescent lights. High frequencies of mating oc-
curred when these cages contained several adults. In these same large
cages small Helianthus plants were available for the females to oviposit
upon, following mating.

Life History

Chlosyne lacinia in central Texas is multiple brooded; a generation oc-
about every 35-45 days under field conditions. Adults fly from March
mber with individuals apparently passing the winter as diapausing

ae (there are five larval instars ). Diapause appears to be

ponse because larvae frequently entered diapause in

VoLUME 24, NuMBER 2 17)

the third instar when they were held in the laboratory under light condi-
tions of short day length (i.e. 8-12 hours).

Larvae in central Texas appear to feed primarily on species of the tribe
Heliantheae (Compositae). The primary larval foodplant from March
through July is the sunflower, Helianthus annuus L. It may feed occa-
sionally on H. cucumerifolius Torr. and Gray, Ambrosia trifida var. texana
Scheele, Verbesina virginica L. and Silphium sp. (Kendall, 1959, and per-
sonal communication). We have also found a few broods on Viguiera
dentata (Cav.) Spreng. and Simsia calva (Engelm. and Gray) Gray. As
H. annuus dies out in August, females begin to oviposit on Ximenesia
encelioides Cay. During this period larvae can be found feeding on both
host plants.

Mating usually occurs in midmorning with copulation lasting about
forty-five minutes. Up to 500 eggs may be deposited in one or two clusters
by a female on the underside of a host plant leaf. Counts made of day-old
larvae from 75 collected wild egg masses revealed an average of 139 (Se =
+ 11) hatched eggs per mass, with a range of 22 to 480.

The initial pale greenish-yellow egg color changes to dark brown 24-48
hours before hatching. The pale yellow larvae of a single egg mass hatch
simultaneously. After consuming the empty eggs, they begin feeding
gregariously on a thin silken web in a tightly packed mass on the under-
side of the host plant leaf. When disturbed, first and second instar larvae
frequently will begin a synchronous jerking movement of the body. The
function of the web is not known, but it may possibly help to maintain
colony unity and protect the larvae from some predators and parasites.

When a leaf has been consumed, one or two larvae will move off the leaf
along the stem followed in single file by the rest of the brood. Each larva
lays down a silken thread which is reinforced by the larvae following it. A
distinct silken trail is thus constructed by the group. Upon arrival at a new
leaf, the feeding aggregation is reformed (Bush, 1969).

Third to fifth instar larvae may enter a quiescent period during the
month of August in Texas when the temperature is high and food plants
are scarce. In the laboratory, unfed larvae in the later stages (primarily
third and fourth instar) will contract and remain quiescent for up to two
months or more and resume feeding and complete development when food
is provided. Mr. Roy Kendall has informed us that he has been able to
maintain unfed larvae of C. lacinia for over a year. Thus it appears that
this species is well adapted to the xeric conditions of central Texas.

Usually by the fourth instar, the larvae within a brood begin dispersing
singly or in small groups and may move to adjacent plants, becoming well
scattered by the fifth instar. Dispersal appears to coincide with the ap-

13 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Life table of Chlosyne lacinia reared on sunflower leaves in the labora-
tory at 21° Centigrade.

_ Ist
instar
larvae Incu- Total
per bation Imnstar Instar Instar Instar Instar time
egg period 2 5 Pupa egg—adult
mass (days) (days) (days) (days) (days) (days) (days) (days )
Mean 138.8 9.10 7.8 A5 On flee 12.0 9.0 53.3
Range 99-480 6=15 7-8 425 3255-9) 42227 —

No. of broods U5 Le 6 6 6 6 5) OA

1 Number of individuals.

2 The range of the total developmental time could not be determined with accuracy because the
egg masses used to calculate the incubation period and the broods used to obtain data on the
length of the larval instars were not necessarily the same.

pearance of larval color patterns which usually begin to manifest them-
selves in the third instar and are quite well developed by the fifth. In one
color phase, the ground color is black, in another the ground color is
almost solid orange, and in a third the ground color is black with an orange
dorsal stripe. More detailed descriptions of the immature stages of this
butterfly may be found in Edwards (1893).

The function of dispersal is not clear, though perhaps it is primarily the
result of food competition. It may also serve to reduce the incidence of
disease, parasitism, and predation among individual larvae of the same
brood. This interpretation is supported by the fact that fourth and fifth
larval instars under crowded laboratory rearing conditions were much
more susceptible to disease, which is apparently quiet contagious, than
were the earlier stages under the same conditions. As will be discussed in
more detail later, mature larvae were also more susceptible to parasitism
than earlier stages and predation appeared to be greatest among third to
fifth instar larvae.

Fifth instar Chlosyne larvae frequently wander a considerable distance
from their feeding site prior to pupation, which may occur on sheltered
sites on a wide variety of surfaces such as Helianthus stems, fence posts, or
leaves. The base color of the pupal case changes after 24 hours from pink-
ish-white to ivory or yellow. It is often speckled with highly variable black
markings, producing intergrades between pure white and almost black
pupal cases. Male pupae are slightly smaller than females and within a
given brood emerge approximately 24-36 hours before the females.

Laboratory breeding experience revealed that emerged females must
usually mature for one or two days prior to mating, thus producing a 2 to

; between the sexual maturity of the male, which may mate the
" emergence, and the female. A delay in the emergence and

VoLUME 24, NUMBER 2 139

TABLE 2. Life table of Chlosyne lacinia reared individually on standard labora-
tory diet with ground sunflower leaves at 22°—23° C.

bation ME e MN Me Ete pudgy see atu
Mean (days) IOWGuT Se AST | 654 807 991 7.32 53.67
So 2:99 EIS £97 287 273 244 259 210
Range | 1014 KIO S412 S58 Rol Bay) Gg 7euen
No. of individuals CnC OMS) Blow witogn) ” (og, 7) Sg
died ee Sy AG vo ORT 1 0 1 43
diapause — 0 0 2) 23 4 0 29

mating of females would tend to reduce inbreeding by encouraging dis-
persal among individuals within a brood prior to mating. This is the op-
posite of maturation in Euphydryas editha and Cercyonis species, where
females can and do mate immediately upon emergence but males must
mature at least 24 hours before mating attempts are made (Emmel, 1969).

Two life tables for Chlosyne lacinia are presented in Tables 1 and 2.
Table 1 was compiled from data collected from mass rearings of broods at
21° C on H. annuus leaves. When at least 50% of the larvae had molted the
date was recorded. As all individuals in a brood usually develop syn-
chronously, this instar molting time involved little intrabrood variance.
Those that failed to molt were usually diseased or parasitized.

Table 2 represents the time required for individual larvae to reach the
adult stage. These larvae were selected at random from different broods
and reared in isolation on laboratory media at 22°—23° C and constant light.
The results obtained from the two methods of rearing appear to be in
fairly close agreement. It is apparent, however, that the life cycle takes
considerably longer in the laboratory than under field conditions, though
accurate life tables have not been compiled for wild broods. The principal
reason for faster development in the field may be the higher diurnal tem-
peratures (to 39° C) which increase larval feeding and hence growth rates.

Parasitism and Predation
Parasitism

It is clear the parasitism accounts for a considerable portion of the
mortality occurring from the egg to the adult stage in natural populations.
However, because of the sampling methods employed and the increased
rate of parasitism with larval maturity, it was not possible to obtain an
accurate overall estimate of the rate of parasitism and its effects on natural
populations.

140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 3. Rates of parasitism by the braconids Apanteles lunatus and A. rufo-
coxalis. Larvae were collected at the instar indicated and held in the laboratory
until the parasites emerged. Fourth and fifth instar broods were collected on plants
isolated from other broods to insure that all individuals came from the same brood.

% Larvae
Parasitized.
Broods Larvae % Broods % Larvae per
No. of Para- No. of Para- Para- Para- Parasitized
Stadium Broods sitized. Larvae sitized. sitized sitized Brood
Instar I 6 2, 518 3 OD 0.6 le
Instar II 66 35 7846 359 53.0 4.6 8.2
Instar III 35 26 3490 365 WAL 16.2 25.0
Instar IV 10 6 1411 292, 60.0 AQT 32.0
Instar V 5 4 222, 108 80.0 48.6 ne

The following parasites have been reared from the eggs, larvae, and
pupae of C. lacinia.

Egg: Trichogramma fasciatium (Perkins) (Trichogrammatidae) is the
only parasite thus far reared from the eggs of C. lacinia. Generally all the
eggs in an infested egg mass were parasitized. Ten out of eighty-two egg
masses (12.2% ) were parasitized from collections made at three localities
(Stockdale, Wilson Co.; Czestochowa and Kenedy, Karnes Co.) on June
18, 1968.

Larvae: One species of Braconidae, Apanteles lunatus (Packard) was
found to parasitize all larval stages. A full grown Apanteles larva never
emerged from a host before the host had reached the third instar. Upon
emerging, the parasite spun a bright yellow cocoon on the leaf beside the
moribund host. The frequency of larval parasitism was directly correlated
with the stage of development as indicated in Table 3. Individuals in
broods collected in the first instar and held in the laboratory were rarely
parasitized (0.6%), while approximately 48.6% of all fifth instar larvae
collected in the field were parasitized.

There was some indication that the three larval morphs were differen-
tially parasitized. Orange larvae, for instance, were rarely parasitized,
while black larvae appeared to be more heavily parasitized by Apanteles

than the striped larvae. These situations of selective parasitism are
now under study

Pupae: Three parasite species were reared from a few C. lacinia pupae

/r pupae in the field. The most frequently encountered
parasite was a small yteromalid, Pteromalus archippi Howards, which
‘acinia pupae. Two other Hymenoptera, the
vinnulus (Cresson) and the chalcid Spilo-

!so been reared from pupae of lacinia. Ap-

ly on the pupal stage of this host, while C.

collected is lar

Oviposits in full
ichneumonid Craticl
chalcis phoenic a Burt

parently S. phoenica

VoLUME 24, NuMBER 2 141

vinnulus was reared from a pupa which was collected as a fifth instar
larva. Two species of Tachinidae, Eupharocera daripennis (Marquart)
and Siphosturmia melitaeae (Cog.), are common lacinia parasites, but ap-
parently oviposit on fourth and fifth instar larvae. Of 177 larvae collected
at random in the last two instars, 14.1% were parasitized by these flies.

Predation

Predators take a heavy toll of larvae and adults, but no field observa-
tions have yet been made on the predation of the egg and pupal states.
One of the most frequently encountered predators was a metallic-blue
pentatomid bug, Stiretrus anchorago (Fabricius). Most often attacking
third and fourth instar larvae, these bugs would sometimes almost com-
pletely destroy a brood.

Members of the wasp genus Polistes ( Vespidae) are also common preda-
tors of primarily fourth and fifth instar C. lacinia larvae. Two species, P.
annularis (L.) and P. exclamans Viereck have been identified, but others
undoubtedly prey on the larvae. Individuals of both species have been
observed to return repeatedly to the same plant and systematically strip a
sunflower of almost all larvae.

Birds and lizards, both common in the study areas, have never been ob-
served to feed on any stage of this butterfly. Unidentified species of
jumping spiders (Salticidae), on the other hand, have been observed
capturing and feeding on adults and the butterfly has also been found in
the webs of the orb spiders Argiope aurantia Lucas and A. trifasciata (For-
skal) (Argiopidae ).

| Acknowledgments

We would like to express our appreciation for the identification of
parasites to B. D. Burks (Chalcididae, Pteromalidae, and Ichneumonidae ),
R. C. Froeschner (Pentatomidae ), P. M. Marsh (Braconidae ), A. J. Menke
(Vespidae) of the Systematic Entomology Laboratory, U.S.D.A.; D. M.
Wood (Tachinidae) of the Canada Department of Agriculture; and H. J.
Einhard (Tachinidae), College Station, Texas. We would also like to
thank N. Price, J. Lee, P. Barton, and R. Neck for their assistance in rear-
ing the larvae. Particular thanks are due Mr. B. W. Records who carried
out the individual rearing test. Roy O. Kendall, San Antonio, Texas, gave
generously of his time and advice on this research.

This work was carried out with support from Grant GM-15769 from the
National Institutes of Health.

Summary

1. The nymphalid butterfly, Chlosyne lacinia Geyer, ranges from South America to
the southwestern United States and is polymorphic in the larval, pupal, and adult

142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

stages. Features of this species’ biology make it exceptionally suitable for studies in
ecological genetics of natural populations.

2. Techniques for rearing Chlosyne lacinia in the laboratory are described. The most
satisfactory method of rearing is on an artificial medium (the Special Vanderzant-Ad-
kisson Modified Wheat Germ Diet, with antibiotics and powdered Helianthus sun-
flower leaf additives) under a 16 hour photoperiod at about 25° C. Adults are kept in
one-pint ice cream cartons with netting tops, and are fed daily with a honey (or sugar)
water solution. Mating will take place in cages under florescent lights as well as sun-
light.

3. The primary natural food plant from March through July in Texas is the sun-
flower, Helianthus annuus 1. When this species dies in August, female butterflies
begin to oviposit on Ximenesia encelioides Cav. Several other representatives of the
Heliantheae (Compositae ) which serve as secondary hosts are discussed.

4. Chlosyne lacinia lays large clusters of eggs and the larvae are gregarious until the
fourth instar when they disperse. A silken trail is laid down and followed by larvae
moving from one leaf to another during the gregarious phase. Black, orange, and
black-with-orange-stripe color morphs occur in the larvae (clearly recognizable at
fourth and fifth instar). Diapause occurs in the third instar, and may occur under nat-
ural conditions of food deprivation (in August) or on the approach of winter (in late
November). Laboratory experiments indicate the winter diapause may be triggered by
short-day photoperiod.

5. Pupation occurs at some distance from the host plant on which the larva fed. The
pupae vary in color from pure white to almost black.

6. Newly emerged males are ready to mate the same day of emergence, but females
must mature for one or two days prior to mating.

7. Parasites and predatoxs discovered to date are discussed.

Literature Cited

Apkisson, P. L., ERMA S. VANDERZANT, D. L. BULL, AND W. E. Auuison. 1960. A
wheat germ medium for rearing the pink bollworm. J. Econ. Entomol. 53: 759-
eee

Busu, G. L. 1969. Trail laying by the larvae of Chlosyne lacinia (Lepidoptera,
Nymphalidae). Ann. Ent. Soc. Amer. 62: 674-675.

Epwarps, W. H. 1893. Notes on a polymorphic butterfly, Synchloe lacinia, Geyer
(in Hub. Zutr.), with description of its preparatory stages. Can: Ent. 25: 286—
291.

uriicH, PAuL R. AND ANNE H. Enriicu. 1961. How to know the butterflies.
Wm. C. Brown Company, Dubuque, Iowa. 262 pp.

EMMEL, THoMAs C, 1969. Taxonomy, distribution and biology of the genus Cercy-
onis (Satyridae). I. Characteristics of the genus. J. Lepid. Soc. 23: 165-175.

et Roy O. 1959. More larval foodplants from Texas. J. Lepid. Soc. 13: 221—

KLots, A. B. 1951. A field guide to the butterflies. The Riverside Press, Cambridge,
Massachusetts. 349 pp.

VoLUME 24, NuMBER 2 143

NOTES ON LETHE CREOLA (SATYRIDAE), WITH
DESIGNATION OF LECTOTYPE

Roperick R. [Rw1n!
24 East 99th Place, Chicago, Illinois

Despite its occurrence in populous and long-settled parts of the United
States, Lethe creola (Skinner ) remains a comparatively obscure and little-
known butterfly, poorly represented in most collections. Several factors
may share responsibility for this. L. creola is very local, and of crepuscu-
lar and retiring habits, so that it may well be overlooked where it occurs.
There has been much confusion of creola with its congener L. portlandia
(Fabr.) which it closely resembles in the female sex. In fact, Gillham
and Ehrlich (1954) found that the female paratype of creola was itself
actually portlandia. The conflicting and erroneous records of the species
and descriptions of its range which have appeared in the literature have
contributed their share to the problems which surround it. Further, the
lack of an adequate figure of the female creola in the popular literature
has probably resulted in the inability of many collectors to recognize it.

During the preparation of a forthcoming checklist of the butterflies of
Illinois, it became desirable to verify the reported occurrence of L. creola
in that state. This investigation quickly developed into a much more ex-
tensive study of the species as a whole, taking into account its taxonomy,
characters and distribution. It is hoped that this paper may clear up some
of the problems above outlined.

Taxonomy

Skinner (1897) described Debis creola from specimens sent to him “by
Mr. G. R. Pilate, who captured them at Opelousas, Louisiana, on July 3rd,
present year.” The number of specimens comprising the type series is not
stated, nor are holotype, aliotype or paratypes designated. In 1926, how-
ever, Skinner referred to the “type” and “allotype” of creola as being the
only examples of the species which the had actually seen, thus clearly
indicating that they were the only specimens before him when he wrote
his description of creola. Mr. Harry Clench, with whom I have discussed
the taxonomy of creola at length, believes that Skinner's statement (1926),
in conjunction with the labels of these two specimens, might be considered
a lectotype designation. He suggested the propriety, however, of making
a formal selection of a lectotype for creola in order to remove any ambigu-
ity which might still exist from Skinner's actions.

1 Honorary Curator of Lepidoptera, Illinois State Museum.

144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The two members of the type series are in the Skinner Collection in the
Carnegie Museum. These are a male, labelled “type” but which has not
been so designated in accordance with the Code, and a female, which is
actually a specimen of L. portlandia. J designate the male specimen,
labelled “Skinner” (white paper, letterpress; “Opelousas/7/3/97 La”
(white paper, letterpress; date in black pen); “TYPE NO. 7039/Debis/
creola/Henry Skinner” (red card; letterpress with species name in black
pen) as the lectotype of Debis creola Skinner. I have placed an appro-
priate label, reading “LECTOTYPE/Debis creola/¢ Skinner/Designated
by/R.R. Irwin 69” (red card; black pen) on the pin of this specimen.

The taxonomy of this species is complicated by the fact that Skinner's
female “type” of creola has been found by Gillham and Ehrlich (1954)
to be a female of L. portlandia. Mather and Mather (1958) discuss this
finding in detail.

In addition to the type specimens, the two examples which Holland
used as models for his figures of creola in the “Butterfly Book” are in the
Carmegie Museum. They are apparently members of the “type lot,” but
cannot be considered syntypes since there is no evidence that they were
ever before Skinner, most probably having been transmitted directly by
Pilate to Holland. It may be assumed, in fact, from Skinner (1926) that
he saw only the illustrations of Holland’s specimens. The case of these
two specimens is analogous to that of the types with respect to specific
identity. Clench (in litt.) has found that the specimen which Holland
used to illustrate the female of creola is, like the female paratype, actually
L. portlandia.

Characters

The recognition of male creola presents no difficulty; the apically pro-
duced forewing and patches of dark, raised androconial scales between
the veins are unmistakable. It is in the opposite sex that identification
problems arise. In wing shape and other respects, female creola strongly
resembles the female of typical L. portlandia portlandia, the subspecies
of the southeastern states. Most authors mention characters based upon
the relative size and number of the ocelli on the ventral forewing. A
study of long series of both species has convinced me that these are of
little value. They hold true for the majority of examples of both species,
but enough individual variation occurs to reduce sharply the usefulness

of such characters. For example, I have seen specimens of L. portlandia
anthedon (Clark) exhibiting five well-developed ocelli, the number also
ed by creola; in these, the ocellus below vein Cus was fully equal

ment to the others. Forbes (1960) ignores characters based on
id + cntions only the single character which appears to hold

VoLUME 24, NuMBER 2 145

Shor

a
| THis se.

FUMEN tS
Paotun,
P Rare THe S ..

Ss.

Figs. 1-5. 1, Lectotype of Debis creola Skinner, upper side; 2, same, lower side;
3, @ paratype of Debis creola Skinner, upper side (actually a @ of L. portlandia; see
text); 4, same, lower side; 5, labels of the above specimens. Those above the wavy
line are of the lectotype; those below it, of the paratype.

All photos by Mr. Allan Watson, Department of Entomology, British Museum
(Natural History); specimens in Carnegie Museum collection.

constant in the female sex: the shape of the postmedian line on the under-
side of the primaries. In creola, this line is irregular and protrudes
strongly outward in cell M,, while in portlandia it is relatively straight.
This character is even more pronounced in the female than in the male,
as shown by comparison of the underside of the male (Fig. 2) with that

of the female (Fig. 7).

146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 6, 7. 6, Lethe portlandia portlandia (Fabr.) 9 New Berne, N. C., leg. S.
Strecker (Strecker Collection), lower side; 7, L. creola (Skinner) 9 Stewart County,
Georgia (Strecker Collection), lower side.

Arrows locate the postmedian line which is the most reliable character for the sepa-
ration of portlandia and creola in the female sex; see text. (Photos by Field Museum
of Natural History. )

Distribution

This study was inspired not only by the need to verify the occurrence
of creola in Mlinois, but by the memory of my years of fruitless search for
the butterfly in well-collected areas from which it had been reported,
principally Palos Park, [linois, and southern Michigan. The results of my
investigation indicate that not only is the actual range of creola consider-
ably less extensive than most present-day butterfly manuals suggest, but
that it is very probably limited by host plant association. It is apparent,
too, that it possesses a continuous range, rather than a disjunct one as
indicated by published reports. The range of creola given by Klots (1951)
is typical of these: “Manitoba, Illinois, Michigan, eastern Virginia s. to
Texas, Louisiana, Florida (rare).” Again, Forbes (1960) seems to have
had a more nearly correct concept of its distribution when he described
it as “In the Mississippi Valley north to Illinois, in the east not seen be-
yond the Dismal Swamp in Virginia.”

While so far as I have been able to determine, the life history of creola
t been worked out, all available evidence indicates that the host
plants are species of Cane (Arundinaria), particularly Switch Cane (A.
tecta ). Canes are restricted to the southeastern and south central portions
of the United States, south of approximate latitude 39°, and west to
ma, and possibly extreme southeastern Kansas.
Kecords of ereola from outside this area were critically examined; most
erroneous. These include the widely quoted records
from Manitoba, Michigan, and Palos Park, Illinois. (The Illinois state

“ . : . , ho : . _ iE - - a .
record m retained, however, as recently true creola has turned up in

has ho

eastern ‘Texa Oklahe

were found to be

VoLUME 24, NuMBER 2 147

én

{ ae "
\ A
Aker
KL/

LL9v

To copy or reproduce this copyrighted map, or any portion thereof, by any method, including office
copying machines, for personal or company use or resale without written permission is illegal.

Fig. 8. The distribution of Lethe creola as compared to that of its putative food-
plant, Arundinaria species. Shaded area represents the range of Arundinaria in the
United States; the dots represent localities from which Lethe creola has been authenti-
cally recorded. In Virginia, each dot represents a county from which creola has been
recorded; elsewhere, each separate locality is shown, except for additional records re-
ceived too late for inclusion on the map; see text.

several localities in extreme southern Illinois.) Records from Kansas
(Randolph, 1929) and Florida (summarized by Kimball, 1965) remain
in doubt because it was not possible to locate and verify the identity of all
specimens referred to in these papers.

Dr. C. L. Remington (in litt.) makes the following very interesting
statement, which does much to explain the confusion of creola and port-
landia: “There is a general myth around that far-southern specimens
which look quite different from the usual more northern portlandia must
be creola. The great Louisiana collector for many years was F. R.
Arnhold, and he was under this misunderstanding and sent large numbers
of Louisiana portlandia to collectors all over the country under the creola
label. We in fact have many of these in the Arnhold collection, now at
Yale, and my father [P. Sheldon Remington] had some.” In view of
Remington’s statement, therefore, all Arnhold “creola” from Louisiana
ought to be viewed with suspicion, and their determinations rechecked

OURNAL OF THE LEPIDOPTERISTS SOCIETY
4i

by whomever may possess them in their collections. Ross and Lambre-
mont (1954) mention these Arnhold “creola” from Louisiana, which are
now known to be portlandia. The types and the Holland specimens re-
mained the only Louisiana records until recently, when Mr. Gayle T.
Strickland of Baton Rouge took several specimens in West Feliciana and
East Baton Rouge parishes. His records, unfortunately received too late
for inclusion on the distribution map, were authenticated by Messrs.
Richard Heitzman and Bryant Mather.

With the elimination of the definitely and probably erroneous records
of creola discussed above, all other localities known to me for the species
are seen to fall well within the range of the species of Arundinaria. These
are indicated in the map (Fig. 8), and are based either upon material
which I have examined, or records which I consider entirely reliable.
These localities are summarized below. In addition to Strickland’s Louisi-
ana records, that from Fayetteville, in northwestern Arkansas, was re-
ceived too late for inclusion on the distribution map. Heitzman and Dr.
Leo J. Paulissen have just informed me (in litt.) that creola is well estab-
lished near that city.

The stronghold of the species would appear to be the Dismal Swamp
area of southeastern Virginia (Clark and Clark, 1939; 1951), but positive
differentiation of the species from portlandia may well prove it to be
commoner elsewhere in its range than previously suspected. This range
may be defined as follows: from eastern Virginia and southern Illinois
south and west, to and including eastern Texas, Louisiana, Arkansas and
southern Missouri. The species is thus seen to be of Lower Austral affini-
ties, as is its probable host plant, and its occurrence beyond the limits of
that subregion, or outside the range of Arundinaria, must be regarded as
unlikely or accidental.

Summary of Distributional Records for Lethe creola

[ list below, by state and county, all localities known to me from which
Lethe creola has been authentically recorded. All are shown on the map
(Fig. 8), with the exception of those marked *. Virginia records are indi-
cated on the map by county only because of the number and closeness of

localities in that state. Abbreviations used are as follows: AMNH =
\merican Museum of Natural History; FMB =F. M. Brown; CNC =
Canadian National Collection; RLC = R. L. Chermock; AHC = A. H.
Clark; CFdP = C. F. dos Passos; DE = Donald Eff; FMNH = Field Mu-

Natural History; FDPI = Florida Division of Plant Industry;
\. Freeman; LH = Lucien Harris, Jr.; RH = Richard Heitz-
linois Natural History Survey; MCN = M. C. Nielsen; LJP

VoLUME 24, NuMBER 2 149

=L. J. Paulissen; GWR=G. W. Rawson; SIU = Southern Illinois Uni-
versity; GTS = Gayle T. Strickland; USNM = United States National Mu-
seum; Yale = Yale University.

ALABAMA: Mobile, Mobile County; Tuscaloosa, Tuscaloosa County
(RLC).

ARKANSAS: Magnolia, Columbia County (RH); North Little Rock,
Pulaski County (HAF); west of Little Rock, Pulaski County (LJP);
Fayetteville, Washington County (RH)*.

GEORGIA: Savannah, Chatham County (DE); Athens, Clarke County;
Stone Mountain, De Kalb County; Atlanta, Fulton County (LH); Stewart
County (Strecker Collection, FMNH); Yonah Mountain, White County
(LH).

ILLINOIS: Benton, Franklin County; Murphysboro, Jackson County;
Lusk Creek near Eddyville, Pope County; Stonefort, Saline County
(SIU ); Pine Hills, Union County (INHS, SIU).

INDIANA: Patoka River near Valeene, Orange County (Masters and
Masters, 1969 ).

LOUISIANA: Baton Rouge, East Baton Rouge Parish (GTS)*; Opelousas,
St. Landry Parish (type locality); Weyanoke, Retreat, West Feliciana
Parish (GTS )*.

MISSISSIPPI: Clinton, Brownsville, Hinds County; Natchez Trace Cy-
press Swamp, Madison County (Mather and Mather, 1958 ).

MISSOURI: Hollister, Taney County (Kite, 1934).

NORTH CAROLINA: Gastonia, Gaston County (RLC, USNM ); Fontana
Village, Graham County (MCN); Hendersonville, Henderson County
(FDPI); Southern Pines, Moore County (FMB); Wilkesboro, Wilkes
County (AMNH).

SOUTH CAROLINA: Greenville, Greenville County (AMNH); Coosa-
whatchie, Jasper County (CFdP, AMNH, Yale).

TEXAS: “Texas” collected by L. Heiligbrodt (Strecker Collection,
FMNH). This may be the “aberration a” of L. portlandia described by
Strecker (1878); probably from Bastrop, Bastrop County, where Heilig-
brodt lived and collected (Brown, 1964).

VIRGINIA: Bog near Petersburg, Dinwiddie County (AHC); near Em-
poria, Greensville County (AHC); Zuni, Isle of Wight County (AHC);
near Adam’s Swamp (AHC), Dismal Swamp near Suffolk (AHC,
AMNH, CNC, CFdP ), Nansemond Escarpment (AHC), Suffolk (GWR,
CFdP, FMB, FMNH), and Raly Pocoson (USNM, FMNH), all in Nanse-
mond County; New Bohemia, Prince George County (GWR, AHC):
Creeds, Princess Anne County (AHC).

150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Acknowledgments

A great many persons gave me assistance in my efforts to evaluate the
authenticity of the various published records of L. creola; space does not
allow me to list them individually, but to all of them I am very grateful.
For their assistance in providing the illustrations, I am indebted to Dr.
Rupert Wenzel and Mr. Michael Prokop of the Field Museum of Natural
History; to Mr. Harry Clench of the Carnegie Museum, and to Mr. Allan
Watson of the British Museum (Natural History). I am additionally grate-
ful to Mr. Clench, to Mr. F. M. Brown, and to Dr. Robert W. Poole of the
Illinois Natural History Survey, for their very helpful criticism of early
drafts of this paper.

Bibliography of Lethe creola

I list below all references to Lethe creola which I have examined in the
course of this study, except mere listings of the species in catalogues and
checklists.

Bancer, F. S. 1958. Euptychia mitchelli (Satyridae) in Michigan and Indiana
tamarack bogs. Lepid. News 12: 41-46.

Brown, F. M. 1964. The types of satyrid butterflies described by William Henry
Edwards. Trans. Amer. Ent. Soc. 90: 323-413.

CuerMock, R. L. 1947. Notes on North American Enodias (Lepidoptera). Ent.
News 58: 29-35.

Criark, A. H. 1935. In Minutes of the 464th Regular Meeting of the Entomological
Society of Washington, April 4, 1935. Proc. Ent. Soc. Washington 37: 115-116.

1936. Notes on the butterflies of the genus Enodia and description of a
new fritillary from Peru. Proc. U. S. Nat. Mus. 83: 251-259.

CiarK, A. H., anp L. F. Ciarx. 1939. Butterflies of a wood road at Suffolk, Va.
Ent. News 50: 1-5.

1951. The butterflies of Virginia. Smithsonian Misc. Coll. 116(7): 1-239.

DE LessE, H. 1957. Revision du genre Lethe. Ann. Ent. Soc. France 125: 75-95.

Dozier, H. L. 1920. An ecological study of hammock and piney woods insects in
Florida. Ann. Ent. Soc. America 13: 325-380.

EBLE P. R., AnD A. H. Enrzticu. 1961. How to know the butterflies. Dubuque,
OWa.

reco, W.D. 1940. A manual of the butterflies and skippers of Kansas (Lepidop-
tera, Rhopalocera). Bull. Univ. Kansas 39: 1-328.

Forbes, W. T. M. 1960. Lepidoptera of New York and neighboring states; Agaris-
lae eee Nymphalidae, including butterflies. Cornell Univ. Agric. Exp. Sta.
Mem. 371.

GILLHAM, N. W., AND P. R. Euruicn. 1954. The butterfly types of Henry Skinner
and co-authors in the Academy of Natural Sciences of Philadelphia (Lepidoptera,

Papilionoidea and Hesperioidea). Trans. Amer. Ent. Soc. 80: 91= ine
HOLLAND, W. J. 1898. The butterfly book. Garden City, New York.
1931. The butterfly book, a new and thoroughly revised edition. Garden
City, New York.
KIMBALL, C. P. 1965. The Lepidoptera of Florida. (Arthropods of Florida, Vol. 1).
Din ision of Plant Industry, Florida Department of Agriculture, Gainesville.
' ake A calendar of Ozark butterflies, Lake Taneycomo region, Missouri.
vs 45: 36-39,

195]

A field guide to the butterflies. Boston.

VoLUME 24, NuMBER 2 15]

Macy, R. W., AND H. H. SHEPARD. 1941. Butterflies. Minneapolis.

Masters, J. H., AnD W. L. MAstTERs. 1969. An annotated list of the butterflies of
Perry County and a contribution to the knowledge of Lepidoptera in Indiana.
The Mid-Continent Lepidoptera Series, No. 6.

MATHER, B., AND K. MATHER. 1958. The butterflies of Mississippi. Tulane Stud.
Zool. 6: 63-109.

Moore, S. 1960. A revised annotated list of the butterflies of Michigan. Occasional
Papers of the Museum of Zoology, University of Michigan, 617: 1-39.

RANDOLPH, V. 1929. A calendar of Kansas butterflies. Ent. News 40:88-92.

Ricuarps, A. G., Jr. 1932. Distributional studies in southeastern Rhopalocera.
Bull. Brooklyn Ent. Soc. 26: 234-255.

Ross, G. N., AND E. N. LAMBREMONT. 1963. An annotated supplement to the state
list of Louisiana butterflies and skippers. Jour. Lepid. Soc. 17: 148-158.

SKINNER, H. 1897. A new species of Debis. Ent. News 8: 236.

1926. Enodia portlandia, andromacha and creola (Lep., Rhopalocera).
Ent. News 37: 42-43.

STRECKER, H. 1878. Butterflies and moths of North America. Reading, Pa.

THE LIFE HISTORY OF EUTRICOPIS NEXILIS
(NOCTUIDAE)

D. F. Harpwick
Entomology Research Institute, Canada Department of Agriculture,
Ottawa, Ont.

Eutricopis nexilis Morrison (1875, p. 102) feeds in the larval stage on
the heads of species of Antennaria, a complex genus of the Compositae,
that is widespread in temperate North America. The insect itself is also
widespread. In Canada, it is distributed from Nova Scotia to south-
central Ontario, and from southern Saskatchewan to southern British
Columbia. There is an apparent hiatus in its distribution through western
Ontario and southern Manitoba. In western North America, the species
is distributed southward at least as far as Colorado. In western Quebec,
nexilis flies during May; with increasing elevation in the west, however,
the period of adult activity is progressively retarded, and at an altitude
of 6,000 feet in southern British Columbia, the flight period may be
protracted into the first part of August.

Eastern and western populations differ in both adult and _ larval
characteristics. Eastern adults are distinguished from western adults by
a less patchy appearance of the dark areas of the forewing and by a
reduction in the extent of the white areas on both fore- and hind wings.
Moreover, representatives of eastern populations are significantly smaller
at the one percent level than their western counterparts, the mean ex-
panse for eastern material being 16.2+1.1 mm! (24 specimens) in

1 Standard deviation

152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

contrast to 18.5+1.1 mm (60 specimens) for western material. The
larvae of eastern specimens are similarly smaller than western larvae,
and most noticeably so in the last stadium. The time required for larvae
to complete their growth is also significantly shorter among eastern
individuals than among western individuals. The mean duration of the
feeding phase of the larval stage of specimens reared from Hull, Quebec
was 15.0 = 0.9 days (18 larvae) and from Keremeos, B.C. 21.3 = 2.9 days
(12 larvae). The two are significantly different at the one percent level.

On the basis of differences in both larvae and adults, eastern and
western populations of Eutricopis nexilis might be considered as repre-
senting different subspecies.

Behaviour

Eutricopis nexilis is an exclusively day-flying species, and has the
greatly reduced compound eyes that are commonly associated with this
period of activity. The little moth flies very quickly and the patchy
dark and white pattern of both fore- and hind-wings makes it almost
impossible to keep the insect in view. In the cooler parts of the day and
during cloudy periods, adults of the species commonly rest on the flower-
ing heads of the food plant.

The eggs are deposited singly among the florets of Antennaria, the
female forcing her abdomen down through the head from the top to
oviposit above the developing seeds. For a noctuid moth of such small
size, E. nexilis deposits an extraordinarily large egg. The pattern of
depositing such a large egg is one that has evidently developed within
some of the smallest heliothidine moths to insure the survival of their
progeny (for discussion, see Hardwick 1969). The fecundity is corre-
spondingly low; five wild-caught females, taken at Hull, Quebec, de-
posited a mean of 7.8 eggs, and the maximum laid by any individual
female was ten. A majority of the eggs observed, hatched on the fourth
day after deposition.

The newly hatched larva feeds first on the florets, then on the devel-
oping seeds. First- and second-stadium larvae show very characteristic
external evidence of their feeding within a head. One to several florets
are severed at their junctures with the seeds and the florets are pushed
outward so that they protrude above the general surface of the head.
Usually during the third stadium, the larva quits the first head and enters
5 iy mal] ing its ay downward through the top of the head toward
the seeds,

Third- and fourth-stadium larvae also give typical external evidence
of their presence within a head. All of the florets are severed from the

VoLUME 24, NuMBER 2 153

Figs. 1, 2. Eutricopis nexilis Morr. 1, Nickel Plate, Hedley, B.C., 6,500 feet:
2, Mt. St. Hilaire, Que.

seeds and pushed outward and a tangled mass of florets and pappi are
left protruding from the receptacle (Fig. 3). The mass does not seem
to become dislodged from the plant, however, and it may be secured into
position by strands of silk. The larger larva thus fashions a nest for itself
within the receptacle so that it may remain concealed while feeding on
the seeds.

Larvae mature in four stadia. At the cessation of feeding, the larva
makes its way to the ground and fashions an elongate cocoon or pupal
cell whose walls are formed of granules of soil and bits of organic débris
(Fig. 8). The pupal cell is formed at or immediately below the surface
of the ground. In one case observed, the larva on ceasing to feed, actually
formed a pupal cell among the heads of its food plant (Fig. 4).

So far as is known, Eutricopis nexilis is entirely single-brooded through-
out its range. It overwinters as a pupa in the earthen cell at the surface
of the ground.

Descriptions of Stages

Data on the immature stages of Eutricopis nexilis are based on
rearings from three localities: Aylmer, Quebec; Twin Lakes, Keremeos,
B.C., 4,700 feet; and Osoyoos, B.C., 4,000 feet. The durations of stadia
listed are those obtained from rearings maintained at room temperature.
Rearing techniques employed were the same as those outlined by
Hardwick (1958). The estimate of variation, following the means for
various measurements, is the standard deviation.

Adult (Figs. 1, 2). Vestiture of body consisting of black, closely appressed
scales overlaid by elongate, yellowish-grey or olive-grey, hair-like vestiture; overlying
vestiture of abdomen usually not heavy. Forewing olive-grey, marked with pale
yellow or white and suffused with red. Basal space broad, extending outward to
incorporate all of orbicular spot; usually a narrow, pale-yellow or white triangle in
anterior region of basal space marking the proximal margin of orbicular; basal space
variably overlaid and suffused with dark red, most strongly so anteriorly and out-
wardly; red suffusion occasionally so extensive as to obscure any olive-grey colouring.

154 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Leagan aay OS O -
a 4

Figs. 3-8. Eutricopis nexilis Morr., immature stages and damage. 3, Tufting of
florets of Antennaria sp. caused by larval feeding; 4, an unusual cocoon formed in
the head of Antennaria sp.; 5, dorsal aspect of fourth-stadium larvae; 6, right lateral
aspect of fourth-stadium larvae; 7, pupae; 8, pupal cell.

Median space pale yellow or white, very narrow because of enlargement of basal
space; anterior half of median space containing a large, dark, usually rectangular,
reniform spot; pale central area of median space often separated from margins of
wing b dark band along costal margin and another along trailing margin. Trans-
erse posterior line usually smoothly biarcuate. Subterminal line irregular, marked
only by colour change between subterminal and terminal spaces. Subterminal space
shades of red or reddish-brown rarely olive. Terminal space olive-grey. Fringe
usually whit le gr ccasionally concolorous with terminal space. Hind wing
dark blackish-br« nite central spots; central spots often coalescing to
form an ir lar whi nd, Fringe white. Underside of forewing with a
pink a brown b pace in ) orbicular and claviform spots often well defined.
\ white medi

‘arge, dark-brown discal spot. A broad dark-

VoLuME 24, NuMBER 2 155

Figs. 9, 10. Apical abdominal segments of pupa. 9, Ventral; 10, left lateral.

brown, outer-marginal band extending forward from outer angle almost to apex.
A white patch, margined proximally with pink, at apex of wing. Fringe brown at
apex, otherwise white. Underside of hind wing with a brown patch at anal angle
and a brown suffusion along inner margin. Anterior half of wing white or pale
grey variably suffused with pink. Two white central spots or an irregular central
band. Fringe white with a brown basal line.

Expanse: Eastern populations, 16.2 + 1.1 mm (24 specimens); western popula-
tions, 18.5 + 1.1 mm (60 specimens).

Egg. Very pale greenish-yellow when deposited; darkening gradually in the next
three days first to a dark yellow and then to a dull orange. Micropylar half of egg
becoming brown on the third day after deposition. Entire outline of larva visible
through chorion within a few hours prior to hatching.

Dimensions of egg: 0.854 + 0.019 mm x 0.539 + 0.019 mm (3 eggs).

Incubation period: 4.4 + 1.0 days (25 eggs).

First-Stadium Larva. Head black. Prothoracic and suranal shield medium to
dark grey. Trunk pale translucent yellow, often with a strong internal orange suffusion.

Head width: Keremeos, B.C., 0.361 + 0.013 mm (9 larvae); Hull, Que., 0.353 =
0.014 mm (16 larvae).

Duration of stadium: Keremeos, B.C., 3.2 + 1.8 days (12 larvae); Hull, Que.,
3.9 + 0.8 days (18 larvae).

Second-Stadium Larva. Head black. Prothoracic and suranal shields dark brown.
Trunk varying from ashen grey to yellowish grey. Setal bases, thoracic legs and rims
of spiracles dark brown.

Head width: Keremeos, B.C., 0.568 + 0.025 mm (26 larvae); Hull, Que., 0.557 +
0.022 mm (16 larvae).

Duration of stadium: Keremeos, B.C., 3.8+ 0.6 days (12 larvae); Hull, Que.,
2.2. + 0.6 days (18 larvae).

Third-Stadium Larva. Head light to medium brown, variably suffused and
mottled with dark brown; dark suffusion often so prevalent as to obscure any paler
colouring. Prothoracic and suranal shields usually paler than head, brown, mottled
with darker brown; prothoracic shield usually with one or two pairs of pale longi-
tudinal lines. Trunk light to medium grey, usually with a strong greenish tone; a

156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

middorsal and a pair of subdorsal brownish longitudinal bands often evident. Rims
of spiracles brown. Proleg shields concolorous with trunk. Setal bases and thoracic
legs medium to dark brown.

Head width: Keremeos, B.C., 0.924 + 0.037 mm (25 larvae); Hull, Que., 0.846 +
0.051 mm (18 larvae).

Duration of stadium: Keremeos, B.C., 4.7+ 1.0 days (12 larvae); Hull, Que.,
3.4 + 0.8 days (18 larvae).

Fourth-Stadium Larva (Figs. 5, 6). Head pale yellowish-fawn or greenish-
fawn, variably mottled and suffused with chocolate-brown dorsally and in centre of
face; head often only weakly marked with brown. Prothoracic shield fawn marked
with dark brown, usually more heavily marked than head; with a pale-yellow mid-
dorsal line and a pair of paler-yellow sublateral bands. Suranal shield fawn marked
with black. Middorsal band varying from reddish-brown to purplish-red. Subdorsal
area consisting of a dark median band and pale marginal lines, the dorsal marginal
line commonly evanescing; median band of subdorsal area varying from a reddish-
brown, paler than that of middorsal band, to a medium green with a faint pink
suffusion. Supraspiracular area similar in color to median band of subdorsal area;
with an irregular and discontinuous, pale greenish-yellow median line passing through
it. Spiracular band greenish-grey, poorly distinguished from suprapodal area. Supra-
podal and midventral areas concolorous, varying from greyish-green to apple green.
Proleg shields concolorous with suprapodal area. Setal bases and rims of spiracles
dark blackish-brown. Thoracic legs medium brown.

Head width: Keremeos, B.C., 1.482 + 0.073 mm (12 larvae); Hull, Que., 1.354 +
0.076 mm (25 larvae).

Duration of feeding phase of fourth stadium: Keremeos, B.C., 9.7 + 2.6 days
(12 larvae); Hull, Que., 5.4+ 1.2 days (18 larvae).

Duration of prepupal phase of fourth stadium: Keremeos, B.C., 6.2 + 3.2 days
(9 larvae); Hull, Que., 4.7+ 0.7 days (12 larvae).

Pupa (Figs. 7, 9, 10). Uniform light yellowish-brown. Rims of spiracles on
a level with general surface of cuticle. Anterior marginal areas of abdominal segments
5, 6 and 7 only weakly pitted. Apex of wing extending posterior to apex of proboscis.
Apex of tenth abdominal segment dorsoventrally flattened and with a truncated
posterior margin; with a pair of conspicuous, short, stout setae on either side; an
additional, smaller, lateral seta anterior to the conspicuous pair; also a pair of sub-
apical dorsal setae.

Length to posterior margin of fourth abdominal segment: Keremeos, BAG, AL sill Se
0.60 mm (16 pupae); Hull, Que., 4.05 + 0.32 mm (4 pupae).

Acknowledgments

| am grateful to Mr. John E. H. Martin of this Institute for assistance
in the field and for the fine photographs accompanying this paper. I
appreciate the assistance of my associate, Mr. E. W. Rockbume who
measured the immature stages and drew the cremaster area of the pupa.

Literature Cited

Be Taxonomy, life history and habits of the elliptoid-eyed
peci« / (Lepidoptera: Noctuidae) with notes on the Heliothidinae.

/ of Heliolonche carolus (Noctuidae). J. Lepid. Soc.

MoORRISO a

Nat Flic \ ‘N\A

North American Lepidoptera. Ann. Lyceum

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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyr F. pos Passos

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CONTENTS

Clench, H. K. Communal roosting in Colias and Phoebis (Pieridae) _.-

Drummond, B. A., G. L. Bush, and T. C. Emmel. The biology and
laboratory culture of Chlosyne lacinia Geyer (Nymphalidae) . ___-

Freeman, T. N. A new species of Cameraria on bur oak in Manitoba
(Gracillaridae) 0

Hardwick, D. F. The life history of Eutricopis nexilis (Noctuidae )

Irwin, R. R. Notes on Lethe creola (Satyridae), with designation of lecto-
EY POS a NI en

Kolyer, J. M. Variations in the markings of Pieris rapae (Pieridae) induced
during the pupal’ stage 00.0

Lafontaine, J. D. A redescription of Strymon borealis Lafontaine (Ly-
caenidae )

Miller, L. D. Two new Mexican Hesperiidae

Oliver, C. G. The environmental regulation of seasonal dimorphism in
Pieris napi oleracea (Pieridae )

Rozman, R.S. A plastic relaxing box for pinned and papered specimens ___-

Sargent, T. D. and S. A. Hessel. Studies on the Catocala (Noctuidae) of
southern New England. I. Abundance and seasonal occurrence of
the species, 1961-1969

Schmid, F. Considérations sur le male d’Ornithoptera allottei Rothschild
et sur la phylogénie des ornithoptéres

117-120

135-142

86- 87

151-156

143-151

125-134

83— 86

120-124

77- 81

81— 83

105-117

Volume 24 1970 Number 3

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY
Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

3 August 1970

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JOURNAL OF

Tue LEPIDOPTERISTS’ SOCIETY

Volume 24 1970 Number 3

THE MIGRATIONS OF THE PAINTED LADY BUTTERFLY,
VANESSA CARDUI (NYMPHALIDAE), WITH SPECIAL REF-
ERENCE TO NORTH AMERICA

C. B. WILLiAMs
Elm Park Lodge, Selkirk, Scotland

The nymphaline butterfly Vanessa [or Pyrameis] cardui Linn., known
in Britain as the “Painted Lady” or “Thistle” butterfly, and in France as
“La Belle Dame’, is probably the most widely distributed of all butter-
flies. It is found, at some time of the year, in every continent except South
America, where it is rare or absent (see Fig. 1).

In Australia, New Zealand and some of the Pacific Islands there is a
form kershawi which has slight differences in structure and wing mark-
ings, and has sometimes been considered as a distinct species. Apart from
this the Painted Lady is very free from local variation in form or colour.
Specimens from Japan, India, South Africa, Europe and N. America are
indistinguishable. Such widespread uniformity is an indication of the
constant intermingling of populations, and is characteristic of many other
insects which migrate with wide distributions.

In the great land mass of Europe, Asia and Africa, it ranges, according
to the season, from the tropics northwards to within the Arctic Circle and
southwards to the limit of the land. In South Africa this is about latitude
43° S. In southern India and Ceylon about 10° N and in the East Indies
about 10° S. The Australian form reaches nearly 44° S in New Zealand
and Tasmania.

Ecologically the Painted Lady prefers drier and more open country and
in the tropics is more often found on higher and drier land. It is never
common in the wet tropical forest, which may account for its absence in
most of S. America.

Over the greater part of its range in temperate climates it is not a per-
manent resident, and, unlike most of the Nymphalidae, seems unable to
survive cold winters in any stage. As a result, such areas depend for their

158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. The world distribution of the Painted Lady, which is found, at one season
or another, in almost all parts of the world except South America.

Painted Lady population on migratory flights from the more permanent
habitats in the warm-temperate and subtropical climates.

In Western Europe and North Africa, where its occurrence has been
studied for many years, there appears to be very little winter survival
north of the Mediterranean. Each spring there is a northward movement,
sometimes very small, sometimes in immense numbers, spreading from
North Africa, and even from across the desert area further to the south.

In the north the butterflies reach Iceland in some years, and have been
recorded beyond the Arctic Circle in Scandinavia and Finland. The main
immigration into the British Isles occurs about the end of May or June,
but sometimes earlier. The immigrants lay eggs and local-bred butterflies
appear about August. Further south in Europe, there may be two or more
generations before the fall. There is no evidence of any winter survival,
but there is increasing evidence of a return flight to the South in the fall
on a much smaller scale than the northward flight in the spring. This
southward movement is difficult to observe in Britain, but appears to aug-
ment in numbers as it goes south, and quite definite flights have been seen
crossing the Pyrenees in September and October. Still further south I
have seen (in late Sept. 1943) a very extensive southward flight over the
Atlantic between southern Spain and the westward bulge of North Africa.

in eastern and southern Africa the Painted Lady is widespread and
there are records of large numbers making sudden appearances and some-
times of definite directional flights. In Ghana and Nigeria records of
abundance are chiefly in September and October.

In the East, as far as the Caspian Sea, there are many records of north-

VOLUME 24, NUMBER 3 159

erly spring migrations, and a very large flight has been observed crossing
the Shandur Pass in Chitral at 16,000 feet.

Still further east and as far as Japan I have found no direct information
of seasonal movement, but it may well occur without being noticed.

In Australia the form kershawi is uncertain in appearance and has at
times been observed in mass flights similar to those in the northern hemi-
sphere. There is also a possibility that the population in New Zealand may
occasionally be reinforced by flights across the 1200 miles of sea which
separates it from Australia.

There are, in addition to the cases mentioned above, many records of
the butterflies far out to sea. In the North Atlantic there is a remarkable
record of several being seen in 1865 on a steamer about half way be-
tween Iceland and Newfoundland, and another record in 1950 of several
being seen on the tropic line more than half way across the Atlantic be-
tween the Mediterranean and the West Indies. I have also one captured
about 500 miles from land in the Indian Ocean. These are probably in-
dividuals which have been blown out of their normal routes by strong
wind currents.

Food Plants and Life Cycle

The caterpillars of the Painted Lady have been recorded on over a
hundred different food-plants, in many different orders, but chiefly the
Malvaceae, the Compositae and the Leguminosae. A number of plants of
economic importance are included. In subtropical regions of Eurasia they
frequently destroy the leaves of Malva parviflora (Khoubeiza in Egypt),
which is an important vegetable food, and in Eastern Europe ( Yugoslavia )
they can be quite serious pests of maize, lucerne, beans, sunflowers, and
soya-beans. On the other hand, their great preference for thistles in tem-
perate countries, such as the northern U. S. A. has made them welcome
visitors there in grazing country.

Little is known of their seasonal life-cycle in the tropical countries, but
in Europe and North America there appear to be one or two broods during
the period of summer residence. The Painted Lady shows no ability to
enter into a definite resting stage (diapause) or to be able to survive a
cold winter in any stage.

The Seasonal Distribution of the Painted Lady in North America

Summer Distribution: In the summer months Vanessa cardui may be
found breeding in any part of the United States from the Pacific to the
Atlantic. It is more irregular in appearance in the northeastern States; and
is not often recorded in the southeast, though if this is due a real rarity or

160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

MICRAN
EvRore

DOES
occuR HERE

OCCAS IO

Fron

z
A
«x
Zz
2
71)
4
Y)
3

i
t

SA BS En

% y Gabmas fis 3 OCRASIONA Uy

COMMOIV

2 POSSIBLE
OVER-WINTERINS

7

‘ig. 2. The approximate summer and winter distribution of V. cardui in North and

f P ;
, tral nerica,

interes: is not certain. It may here be confused with V. virginien-
nerally more common.

wn to breed in the southern portions of all the prov-

i nbia to Newfoundland. In the latter it is sporadi-

VOLUME 24, NUMBER 3 ILL

cally quite common. In the extreme north, adults have heen seen on the
Labrador coast, and in the Northwest Territories at Chesterfield Inlet on
Hudson Bay, in north latitude 64°. This is nearly as far north as Ice-
land to which the migrant Painted Ladies from the European fauna oc-
casionally stray. Adults have also been captured on Great Slave River in
the Mackenzie District at about 60°30’ N.

I have no records from northern British Columbia or from the Yukon,
although I would have expected it to go at least as far north in the west
as in the east, particularly as the summer isotherms run farther north in
the west. The only evidence for its occurrence in Alaska is that W. H.
Edwards in 1887 mentions it, without any details, in a list of butterflies
known to occur there.

The Painted Lady also appears sporadically in numbers in the Bermuda
Islands, about 600 miles east of South Carolina. The origin of these may
be in the Antilles, to the south, but this is not certain.

Winter Distribution: In the winter months, by contrast, there is no
evidence of survival in any stage in any part of Canada or the United
States, except possibly in Florida, along the Gulf Coast, and more def-
initely in the extreme southwest of California, where breeding has been
recorded in mid-winter.

In Mexico V. cardui appears to breed throughout the year, sometimes in
abundance, but more records, including the months of activity are still
needed. Still further south in the Central American States it becomes less
common as the climate changes to the wet-tropical type. In Panama it is
rare and in S. America almost unknown.

It occurs at times, and may breed, in most or all of the Caribbean Is-
lands, but not on any scale of abundance that could account for the
masses of the northerly spring migrations. In the most southerly island of
the Antilles, Trinidad, it is a rarity. I know this from my own experience,
as I lived in Trinidad for six years, and did not see one.

In the Hawaiian Islands in 20° N latitude, the same as that of Southern
Mexico, one of the earliest records was reported by Blackburn in 1879.
He stated that it was abundant in that year, though he knew of no previous
record and had seen none before in his three years residence. Today it
appears to be accepted as a regular breeding resident.

Pattern of Migration

The only solution that appears to fit our present knowledge is that the
main, and perhaps the only, source of the parent swarms which invade N
America in the spring is to be found in those parts of Mexico, particular]
in the northwest, which have a relatively dry climate. This area cor-

162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

responds both in latitude and climate with the semi-desert parts of N.
Africa which provide the main source of our European immigrants.

This would imply that the immigrants that appear fairly regularly in
Newfoundland, or their parents, have their origin in western Mexico,
nearly three thousand miles away.

Such mass movements could be carried out either by large numbers
being blown by strong winds far out of sight at high altitudes, or by
directional movements near the ground, possibly independent of the
wind. For the former type of distribution we have at present no evidence,
but this is not proof that it does not occur. For extensive movements at
ground level evidence is steadily accumulating and a summary is given
below.

Direct Evidence of Northward Spring Migration at Ground Level

Over the course of many years I have collected nearly 100 records of
flights seen by naturalists in North America when the butterflies were all
moving definitely in one direction. These vary from observations of a
few dozen butterflies only, to one (quoted below) which was estimated
to include 3000 million individuals. The characteristic, as in other migrant
species and in other parts of the world, is a continued “purposeful” flight
in a steady compass direction, apparently independent of the angle of the
sun or the direction of the wind (see later). Such flights may last an hour
or two, or go on for days and, though usually observed at a single locality,
may cover a front of a hundred miles or more.

It might be interesting to quote here from E. A. McGregor’s account
(1924) of a gigantic movement of Painted Ladies to the north in Cali-
fornia, to give an idea of a migration in its extreme form. He writes:
“Continuing from 11th to 13th inclusive, there was a remarkable migra-
tion of this butterfly. All through the day there was an average of about
300 butterflies per acre at a given moment. The flight direction appeared
to be from south-east to north-west.

“In travelling, the flight was not characteristic of butterflies in general,
but was of a more steady and purposeful nature. During calm intervals
the flight took place on an average of perhaps 10-20 feet altitude, but
during the periods of windiness the butterflies flew very close to the
ground. ‘There appeared to be no attempt at pairing and the individuals
parated—possibly 10 feet apart on the average. It was very

| | pursued their flight in the same direction. It was
‘5 were seen to alight on vegetation but this they

irred at the warmest period yet experienced this

VOLUME 24, NUMBER 3 1163

27>7%D
P°)
a

$5 nw

A
~
A
NN
A
A
nN
x
N~
WS
“~
x
A
Za
A
A
A
~
*
IN
A

AUGUST
AND

Fig. 3. Analysis of the direction of flight in nearly 100 recorded flights of migrat-
ing Painted Lady butterflies in North America, divided into those which occurred in
the spring and early summer up to the end of July; and those which occurred in
August and the fall. The records provide strong evidence for a large northerly flight
in the spring, and a smaller return to the south in the fall.

season, temperature maximums ranging from 80-88° F. The temperature
suddenly dropped late on the afternoon of the 13th, accompanied by a
chilly wind, and the flight as suddenly terminated. On the 15th the
temperature rose again, but a gusty wind occurred, so that only a few
individuals could be seen migrating.

“An attempt to estimate the number of individuals comprising this
three-day migration is here presented. The flight was estimated to

164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

I 2

TaN
NN
I~
NN
NN
wn
NN
AN
Nn
IN
IN
IN
TAN
*
ran
Aa

Fig. 4. The records of flights for the spring months, as detailed in Fig. 3, sorted
according to the state or province in which they were seen.

have been at least forty miles in width (it no doubt was much wider).
The daily duration of the migration was at least twelve hours, or a total
for three days of thirty-six hours flight on any given point.

“Now the rate of travel was estimated at 12 miles per hour, which would
give a total dispersion length of 432 miles for the three days. Within such
a zone (40 miles by 432 long) is contained about 17,280 square miles or
| 1,059,200 acres. With the established occurrence of about 300 butterflies
per acre, it is readily computed that at least 3,000,000,000 had passed on
the 40-mile wide zone between Sunland and Lemon Cove.”
eturn to the evidence available for the northerly spring flight,

ho | summary of the direction of movement in 96 recorded
nave been able to trace, arranged according to the time of

54 occurred before the end of July (mostly from April to

June) a | ‘ve an overwhelming trend to the NW, N, and NE
quarter, | hows the same flights sorted according to the state or

VOLUME 24, NUMBER 3 165

province, and it will be seen that the NE component is almost completely
accounted for by records west of the Rockies from California and Arizona.
Records of spring flights in the middle and NE states are very few, and
are completely lacking from the SE.

This northerly flight, at or near ground level in North America, has its
exact counterpart in Europe, where we have hundreds of observations of
such movements from the Mediterranean northwards; and also smaller
numbers from northern Africa, and others at sea crossing the one to four
hundred mile barrier of the Mediterranean Sea, or coming in to southern
England across the Channel.

Evidence for a Southward Migration in the Fall

If North America is repopulated each year from the south, do all the
offspring of the earlier immigrants die out before the fall, or is there a
partial or general return to the south, to repopulate or replenish their winter
headquarters?

Fig. 3B shows the very little evidence from North America that we have
available on this point. There are only a dozen observations of flights after
the beginning of August, and all but one of these have a southerly com-
ponent, and eight of the twelve were said to be flying “to the south”. The
states and provinces, in which these flights were seen, are California (2),
Colorado, Minnesota (several years, but no definite dates), New Jersey,
New York, Florida, Ontario and Manitoba.

One of the most definite of these flights was seen by W. B. Henson at
Sault Ste. Marie in Southern Ontario on the 30th Sept. 1949, and several
days following. He estimated that 100 to 150 butterflies were crossing a
ten-yard front every ten minutes over a front of at least a mile. This is
equivalent to approximately 100,000 butterflies per hour on the observed
mile. All the butterflies were flying to the south across a westerly wind.

It will be noted that many of these fall records are from the more
easterly states, where mass flights in the spring are not so regularly
recorded. This may perhaps be a psychological effect in that observers in
the west are so satiated with Painted Ladies by the time the fall comes,
that they cease to be interested.

The flight in Florida, referred to above, was that two V. cardui were
captured by Mr. and Mrs. Hodges, along with several V. virginiensis in a
long continued movement to the south of many species of butterflies in
the fall of 1952. Further observations in this area are much needed.

As with the spring flight, the probability of a fall movement to the
south in North America is supported by many records of a similar move-
ment in Europe. My own observations of thousands of Painted Ladies

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

flying to the south far out at sea between Spain and the bulge of Africa in
September 1943 has already been mentioned. The butterflies were very
far apart, but the flight covered the whole fifteen hundred miles from off
the mouth of the Mediterranean to the latitude of Sierra Leone, and was
observed at all times of the day for eight successive days, and mostly fifty
to a hundred miles from land.

Year to Year Variation in Abundance and Range

In some years the immigrant Painted Ladies and their larvae are
abundant over most of the United States and southern Canada; in other
years they may be so rare as to escape notice except in a few places. Such
variations in numbers almost certainly reflect different weather conditions
either in their winter breeding areas, or in the districts they pass through
on their way north.

In the course of years I have collected between one and two thousand
records of their presence (rare, common or abundant) in N. America in
different years and in different areas. Table 1 shows a very condensed
summary for the years 1900-1959, indicating when the immigration has
been above average. It must always be remembered that “absence of
evidence” is not the same as “evidence of absence”; and the further one
goes back, the more frequently one is frustrated by the former state of af-
fairs. In general there are more records of mass flights and unusual abun-
dance west of the Rockies. There are however only a few records of either
abundance or migration in Texas, which suggests that the immigration
from Mexico is more concentrated in the western half of the country.

Figs. 5 and 6 show the constrasting conditions in the two consecutive
years 1951 and 1952. In the former year there was scarcely a record of
abundance anywhere, and an unusual number of cases commenting on
its rarity or absence. By contrast, in 1952, the species was abundant in
most of the Pacific States, with records of mass flights through the north-
central and northeastern U. S. and the whole of southern Canada as far
east as Nova Scotia, and as far north as Chesterfield Inlet (64° N) on
Hudson Bay. There are however no records of abundance from Texas or
from any of the Gulf States except Mississippi, where it was said to have

been “abundant”.
Iossihle Relatinn Betrxa0 6 a
Possible Relation Between the Abundance of Immigrant Populations in
AT aAat ) s

estern Europe and North America

> varying abundance of V. cardui from year to year in

NJ a A “aves. } rie t iy 1 1
North Ai riefly in Table 1, makes it possible to compare the

VOLUME 24, NUMBER 3 167

TABLE 1. THE YEARS BETWEEN 1900 AND 1959, IN WHICH, ACCORDING TO
THE AVAILABLE EVIDENCE, THE PAINTED LADY BUTTERFLY WAS MORE THAN
USUALLY COMMON (C) IN NortH AMERICA AND IN BRITAIN

Brit. N. Am. Brit. N. Am. Brita Ne Ams:
1900 Cc c 1920 c Cc 1940 Cc c
Ol — c SAIL c — Al Cc ©
02 Cc Cc YP) 6 - 49, — —
03 Cc — 23 Cc — 43 Cc —
04 Cc Cc 24 — Cc 44 — —
05 = = 95 Cc - 45 Cc Cc
06 c 26 c c 46 Cc 6
07 = Cc 27 6 = 47 6 Cc
08 Cc - 28 Cc - 48 Cc -
09 — — 29 = = 49 c c
1910 = — 1930 — - 1950 6 —
La — — 31 c Cc Sl — —
12 Cc — 32 - = 52 Cc 6
13 Cc c 33 = = 53 = =
14 Cc c 34 — — 54 —
15 = — 35 — Cc 55 c —
16 — — 36 — Cc 56 = -
17 Cc = 37 c = 57 = Cc
18 c 38 c — 58 c 6
19 c c 39 c 6 59 Cc

changes here with those observed in western Europe over the same
period.

In this part of the Old World, interest in the immigration of butterflies
from the south has. been high for many years. In Britain we have annual
reports on migrant insects, including the Painted Lady, for over thirty
years; in Holland, since 1940, in Ireland since 1950, and in Austria since
1956. Also a study of earlier records, from 1850 to 1930, from old natural
history books and journals in Britain, was published by Cockbill in 1942
(Williams et al., 1942). The information available for the years 1900-
1959 for years of unusual abundance in Britain has been added to the
North American records in Table 1. From this double set of data Table 2
has been made, showing in how many years, out of the total of sixty, the
Painted Lady has been common in both continents (A); in how many
years rare in both continents (D), and how often common in one and
rare in the other (B and C).

It is possible to calculate from the total number of “rare” and “common”
years in each continent, how many of the sixty years should be in each of
the four classes, on the assumption of no relation whatever between the
two. These numbers are shown in brackets in Table 2.

168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

RECORDED

faa Re as RARE OR
/ NY, ABSENT

%

{954

B= BREEDING
7 = MIGRATING

(itt = FEW

ONLY *

1952

wn cistribution of immigrant Painted Lady butterflies in
951 and 1952. The former, a year of almost complete
c of widespread immigration.

VOLUME 24, NUMBER 3 169

TABLE 2. AN ANALYSIS OF THE RECORDS IN TABLE 1 TO SHOW THE NUMBER

OF YEARS, OUT OF 60, IN WHICH THE PAINTED LADY was: (A) UNUSUALLY

COMMON IN NorTtH AMERICA AND IN BriTAIN; (B) COMMON IN NoRTH

AMERICA BUT NOT IN BRITAIN; (C) COMMON IN BRITAIN BUT NOT IN N.

AMERICA; AND (D) RARE IN BOTH AREAS. THE NUMBERS iN BRACKETS ARE

THE NUMBER OF YEARS EXPECTED IN EACH CLASS IF THERE WERE NO ASSOCI-
ATION BETWEEN THE TWO AREAS

IN BRITAIN

common rare total

c
2)
E
<
Bato
or
Lu
=
<
a
Ben 2
rif
O
2
=
S
~_
2)
~~

It will be seen that in the classes of “simultaneous abundance” the ob-
served number of years are both greater than that calculated on the as-
sumption of no relation, but in the two classes of “common in one continent
and not in the other”, the observed numbers are below the calculated.

A statistical calculation (using Yate’s correction for small numbers )
gives the parameter 7 as 2.78, which, from 60 observations and one de-
gree of freedom, indicates a probability of 0.10 that the result obtained
could be due to chance. Otherwise the chances are 9-1 the result has
some significant cause. This is not proof of a relation, but suggests a pos-
sibility worth further investigation. It is however interesting to note that
Grant (1937) has already shown a similar relation between high levels of

170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

N WIND

WITH THE WIND

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ALROSS rh ACROSS

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\ 73

SEE: oe ee oe

Pig. 7. (A) N‘ 00 observed flights of the Painted Lady butterfly, from all
parts of the wor ich both flight direction and wind direction were recorded,
arranged acco} 0 the angle between the flight direction and the wind. (B)

seventeen record th America arranged on the same plan.

VOLUME 24, NUMBER 3 fel

immigration of the hawk-moth Celerio lineata Fabr. in North America and
its subspecies C. l. livornica Esp. in Europe.

Since the idea of common origin for the two continents can be ruled out,
it seems likely that there is some common factor, presumably meteorologi-
cal, which bring about the unusual abundances, either in Europe or in
North America, or more probably in the sources of the immigrants in
North Africa or in Mexico. If this is so, such specially favourable con-
ditions cannot be local events, but must include causes which are linked
together from North Africa (10° W to 30° E) to Mexico (100°-120° W
longitude ), a distance of nearly one third of the earth’s circumference.

The Problems of Orientation and Wind

There is no doubt that in North America, as also in Europe and North
Africa, there are very regular movements of insects which are different
in direction at different seasons of the year. They normally have a north-
erly component in the spring and early summer and a southerly com-
ponent in the fall.

The mechanism of this orientation is still uncertain. It has been at-
tributed to the direction of the sun’s rays (polarised or unpolarised ), or
to the earth’s magnetic field; but most often to the direction of the wind.
The main wind-orientation theories are first that the insects are helplessly
blown by the winds and take no part themselves in determining the gen-
eral direction of movement of the population. The second suggestion is
that the insects are able to appreciate the direction of the wind while they
are flying, and can either fly at a constant angle to it or can straighten out
their flight in the direction in which they “want” to go, by allowing for
the drift.

It is of course obvious that when an insect is flying in an air current of
greater velocity than its own maximum speed, the insect must drift down-
wind. The smaller it is, the more likely it is to drift. The main aerial move-
ments of insects like Aphidae and small Diptera must be generally down-
wind. But the more powerful fliers such as the larger butterflies and
moths can fly steadily against moderate winds, and the butterflies usually
shelter during periods of strong wind. The locusts are an exception as,
although they have an air speed of about ten miles per hour, the swarms
often rise to several thousand feet, where the wind speed is greater than
this. As a result these high-flying swarms tend to move down-wind.

I have seen Painted Lady butterflies in Egypt flying, against a light
wind, at a ground speed of about 12 miles per hour; and other observers
have given ground speeds of 15 to 18 miles per hour (the latter also
against a light wind). In California McGregor has also estimated a speed

79 OURNAL OF THE LEPIDOPTERISTS SOCIETY
j WA

of about 12 miles per hour, so that provided that they fly near the ground,
they can make headway, except in high winds, when they tend to settle.

The little evidence we have for North America about the relation of
flight to wind direction is shown in Fig. 7B, in which all the available rec-
ords are shown; the wind is assumed to be blowing from the bottom of
the diagram. The number of records is small, but they show almost equal
numbers of flights with the wind, against it and across to left or to right.
This evidence is very strongly upheld by a large number of records of
flight direction and wind in V. cardui in other parts of the world, which
are summarised in Fig. 7A. There are 473 observations by many different
observers in many different countries and they give no support to the idea
that there is within sight of the ground, any constant relation between
wind direction and flight. The slight excess of flight records with or against
the wind is probably a biassed observational error, as observers are more
likely to take notice of cases where the wind is with the flight (“obviously
the cause” ), or against it (“how curious” ), than merely across.

Association of Species in a Migratory Flight

A remarkable feature of the northward spring migration of the Painted
Lady into Europe is its frequent association with two other species of
Lepidoptera, a noctuid moth Plusia gamma Linn., known popularly as the
“Silver-Y”, and a pyralid Nomophila noctuella Schiff. Many records exist
over the last hundred years of the simultaneous appearance in numbers of
all these species. It has been noted at sea in the middle of the Mediterra-
nean, on the south coast of France, in the British Isles and elsewhere. In
Yugoslavia, where the caterpillars of the Painted Lady destroy soya-beans
and other crops, the association is regular that it has been suggested that
the use of light traps to attract P. gamma might give an early warning of
possible immigrations of the butterfly.

The pyralid known as N. noctuella is widespread in N. America, and
lor many years it was thought that Plusia gamma also occurred, but the
North American species has been separated off under the name of pseudo-
gamma. Little appears to be known of its migratory habits.

In all the available history of outbreaks of V. cardui in America I have
found only one reference to a similar association, when damage by cater-
pillars of V. cardui and Plusia ni Hubn. occurred simultaneously in Utah
in 1965. This species (related to P. gamma) is known as a rare immigrant
into Britain.

Close observations are needed t

0 see if any association of species occurs
in North America

similar to those in Europe.

VOLUME 24, NUMBER 3 173

1958 APRIL \ST PALM SPRINGS tT S.E

Year Month and Dates Locality ARIZONA Flight direction if definite 3

| LiGHTD ON. W. ee

ON (SYard
Time of day Wind direction and force Numbers S.¥A FRONT No. Captured ; in whose collection

FLICKHT THINNED ouT ABOUT
3 MILES Te west

Sex Condition (fresh : worn) Notes :—Extent of flight ; speed ; pairing : egg-laying ; etc.

NEXT Tuo DAYS Coun WET, Wind? 4-7 fT
NONE SEEN SINCE

Present and past weather conditions ; temperature, rain, sun, cloud, thunder, barometer, etc. Height above ground

RALPH DAWSON LETTER OF 20
(4 DESERT LANE. R2 MESA pate

Observer and Recorder (names and addresses) ARI2 ONA Source of information ; reference
Y: carDdu( NONE MENTLONEOD RK.D,

Name of species Other species in flight Identified by

Fig. 8. The standard record card used in Britain for filing information of migrant
insects, filled in from a letter describing a migration of Painted Ladies in Arizona in
1958.

Discussion

My object in writing this brief account of the status of Vanessa cardui
in North America is three-fold: firstly, to summarise our knowledge and
ignorance of the facts, and to emphasise what seems to be their basic
pattern; secondly, to get more information on the history and past out-
breaks to check our conclusion; and thirdly, to get fuller information about
immigrations in the future.

The first of these requires little comment.

For the second, there must be lying about in note books, files, collec-
tions, museums, in obscure journals and in memories, much information
about the occurrence of the Painted Lady in past years, and in different
parts of the continent. Every record which includes at a minimum the year
and the locality, with some idea of the numbers, is a new piece to fit into
the jig-saw picture.

For observations in the future, we can only hope that if entomologists
are prepared in advance they may be able to observe new details, when
they are fortunate enough to witness an actual migration, or even merely
the presence or absence of the butterfly. For many years in Britain we
have been using a standard record card for summarising information. Fig.
8 shows one of these on which is extracted information of a migration of

174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

V. cardui in Arizona. The different sections on the card give some idea of
the points that should be noted and recorded. It is unlikely that all can be
filled in, but as many as possible should be attempted, and particularly the
direction of flight and of the wind (with any variations during the ob-
servations ). If specimens can be captured, dissections of the female to see
if fat body or developed ovaries are dormant, can give important informa-
tion on probable previous or later activity.

The field study of insect migration is a science in which little can be
planned in advance: also observations can seldom be repeated. The
observer must be ready to take advantage of the opportunity of the
moment, and all students must depend on the experience of others as
well as their own.

“By mutual confidence and mutual aid
Great deeds are done, and great discoveries made.”

Please make sure that your observations as well as your conclusions are
made available for others.

Bibliography

Assott, C. H. 1950. Twenty-five years of migration of the Painted Lady butterfly,
Vanessa cardui, in southern California. Pan Pacific Ent. 26: 161-172.

1951. A quantitative study of the migration of the Painted Lady butterfly,

Vanessa cardui L. Ecology 32: 155-171.

-———— 1959. The 1958 migration of the Painted Lady butterfly, Vanessa cardui
(Linnaeus) in California. Pan Pacific Ent. 32: 83-96.

1963. A migration problem—Vanessa cardui, the Painted Lady butterfly. J.

Lepid. Soc. 16: 229-232.

BLackBurn, T. 1879. V. cardui in Hawaii. Ent. Mon. Mag. 16: 161.

Brown, F. M. 1936. Vanessa cardui migrations in the United States. Entomologist
69: 169.

CAMPBELL, R. KE. 1924. (Painted Lady butterfly). Ins. Pest Surv. Bull. 4: 70-71.

Ciark, A. H. 1932. The Butterflies of the District of Columbia. U. S. Nat. Mus. Bull.
toy (ae Be

Comstock, J. A. 1927. Butterfly migration. In Butterflies of California. Los An-
geles, pp. 127-130.

Dawson, R. W. 1937. Records of a butterfly migration (Pyrameis cardui ). Ent. News
48: 248-250.

Dow, R. P. 1924. Migration of Pyrameis cardui. J. New York Ent. Soc. 32: 121.

~ *: (1926. Migration of Pyrameis cardui. J. New York Ent. Soc. 34: 287-288.

IMMEL, ‘I. C. anp Wosus, R. A. 1966. A southward migration of Vanessa cardui in

| late summer and fall 1965. J. Lepid. Soc. 20: 123-124.

I. VERMANN, B. W. 1914. A note on the abundance of the Thistle Butterfly (Pyrameis
cardui). Ent. News 25: 415.

FARNHAM, G. D. 1895. (untitled note) Ent. News 6: 150.

Grant, K. J. 1937, A historical study of the migrations of Celerio lineata lineata
Fab. and C. lineata livornica Esp. Trans. R. Ent. Soc. London 86: 345-357.
Harpy, G. A. 1954. Notes on the occurrence of the Painted Lady, Vanessa cardui, in

the Queen Charlotte Islands in 1952. Proc. Ent. Soc. B. C. 50: 37.

VoLUME 24, NUMBER 3 Hie

1959. Painted Lady, Vanessa cardui, on Vancouver Island. Proc. Ent. Soc.
be G2 56: 30.
Haskin, J. R. 1933. Butterfly clouds, their origin and infrequent occurrence. Ent.
News 44: 255-257.
Howe, W. H. 1967. A migration of Vanessa cardui (Nymphalidae) in Montana
and Wyoming. J. Lepid. Soc. 21: 39-40.
Hoyinec, L. A. 1968. A migration of Vanessa cardui in Ohio. J. Lepid. Soc. 22: 118—
119.
KNOWLTON, G. E. 1954. Migrations of Vanessa cardui, the Painted Lady Butterfly,
through Utah. Lepid. News 8: 17-22.
McGrecor, E. A. 1924. Painted Lady butterfly (Vanessa cardui). Ins. Pest Surv.
Bull. 4: 70.
1926. Painted Lady butterfly (Vanessa cardui). Ins. Pest Surv. Bull. 6:
38.
SAUNDERS, W. 1884. Annual address of the president of the Entomological Society of
Ontario. Can. Ent. 16: 204-213.
ScuppDER, S. 1876. A cosmopolitan butterfly. Amer. Nat. 10: 392-6, 602-11.
SuGDEN, J. W. 1937. Notes on the migratory flights of Vanessa cardui in Utah. Pan
Pacific Ent. 13: 109-110.
SUGDEN, J. W., Wooppury, A. M. AnD GILLETTE, C. 1947. Notes on the migratory
flights of Vanessa cardui in 1945. Pan Pacific Ent. 23: 79-83.
TANNER, V. M. 1941. Painted Lady butterfly in migration. Great Basin Nat. 2: 104.
TILDEN, J. W. 1962. General characteristics of the movements of Vanessa cardui. J.
Res. Lepid. 1: 43-49.
WitpermMutH, V.L. 1924. Painted Lady butterfly (Vanessa cardui). Ins. Pest Surv.
Bull. 4: 128.
WiuiAMs, C. B. 1925. Migrations of the Painted Lady Butterfly. Nature 115: 535-
Bale =
1926. Further records of insect migration. Trans. Ent. Soc. London (1926).
pp. 198-202.
1928. Collected records relating to insect migration. Trans. Ent. Soc. Lon-
don (1928). pp. 79-91.
1930. The Migration of butterflies. Edinburgh.
1933. Further collected records relating to insect migration. Trans. R.
Ent. Soc. London 81: 103-115.
1937. Butterfly travellers. Nat. Geogr. Mag. 71: 568-585.
1938. Recent progress in the study of some North American migrant butter-
flies. Ann. Ent. Soc. America 31: 211-239.
1939. Some records of butterfly migration in America. Proc. R. Ent. Soc.
London (A) 14: 139-144.
1945. Occurrence of Vanessa cardui at sea off the west coast of Africa.
Proc. R. Ent. Soc. London (A) 20: 4—5.
1949. The migration of butterflies in N. America. Lepid. News 3: 17-18.
1958. Insect Migration. London and New York.
WiuiraMs, C. B., CocksiL, G. F., Gress, M. S., AnD Downes, J. A. 1942. Studies in
the migration of Lepidoptera. Trans. R. Ent. Soc. London 92: 101-283.
WeEeEpBuRY, A. M., SUGDEN, J. W., AND GILLETTE, C. 1942. Notes on the migration
of the Painted Lady butterfly in 1941. Pan Pacific Ent. 18: 165-176.

176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

VARIATION OF GRAPHIUM MARCELLUS IN MISSISSIPPI
(PAPILIONIDAE )!

BRYANT MATHER?
913 Mt. Salus Dr., Clinton, Mississippi

Introduction

Graphium marcellus (Cramer) was reported from Mississippi (Mather
and Mather, 1958) as “apparently generally distributed and locally rather
abundant; February through August. The spring form, with shorter tails,
is found from February through April; the summer form, with longer tails,
May through August. The earliest record... is... Ballground, Warren
Co. on 9 February 1957.” A group of 67 Mississippi specimens (48 ¢ ¢,
19 22) taken on dates from 9 February to 27 August, between 1951 and
1967, at 20 localities in 12 counties is now at hand and has been examined.
The 12 counties include three in the northerm part of the state (Chicka-
saw, Tishomingo, and Tunica), five in the central (Claiborne, Copiah,
Hinds, Rankin, and Warren), and four in the southem part (Forrest,
Harrison, Jackson, and Pearl River). A September record: 24 Sept. 1966,
Tupelo, Lee Co., John Bryson, was provided by Charles Bryson; I have
not seen this specimen. The sample of 67 specimens was arranged by date
of capture and data were recorded for sex, forewing length, tail length,
tail pattern, and other parameters. The results are presented and dis-
cussed as they relate to seasonal and other kinds of variation, nomen-
clature, and distribution.

Characters

Sex: The literature on G. marcellus contains few references to charac-
ters that might conveniently be used to separate specimens by sex. Clark
(1935), in describing the kite swallowtails, stated “the abdominal margin
of the hind wings is broadened in the males, and usually bears a distinct
scent organ.” Field (1940) wrote of this group “the inner margins of the

hindwing .. . are turned upward and rolled over, forming pockets that
contain androconia.” Inspection of the inner margins of the hind wings of
the Mississippi specimens revealed differences that appeared to be con-
sistent and the series was sorted by sex on the basis of these differences.

0 pl : Wee .
"he specimens were subsequently examined by Dr. John M. Burns on 19

Contr bution No. 143, Bureau of Entomology, Division of Plant Industry, Florida Department of
culture, Gainesville, Fla.
h Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida

Agriculture,

VOLUME 24, NUMBER 3 a

FOREWING LENGTH, MM

SPRING SUMMER

Oy A O
eo A ®@

lo Ww Weems =O. 22 947 26 . 28) 2°
TAIL LENGTH, MM

Fig. 1. Relation of forewing length to tail length for 59 specimens of G. marcellus
from Mississippi, showing division of the group into spring and summer subgroups
and boundaries of available data.

August 1967 and I was pleased to find that my sorting of this series by
sex was confirmed by him in each case. I suggest that future writers note
this diagnostic criterion for the guidance of others.

Dimensions: Each specimen on which such measurements could be
made (eight of the specimens had neither tail complete) was measured
to the nearest millimeter for forewing length and tail length. The range
in forewing lengths of the 67 specimens is from 30 to 47 mm. The range in
tail lengths of the 59 specimens is from 11 to 29 mm. The forewing and
tail lengths are plotted in Fig. 1. On Fig. 1 are also shown lines of loci
of forewing length to tail length ratio of 1.5:1, 2:1, and 3:1. Since the
forewing length : tail length ratio changes with change in forewing
length and tail length, the ratio was calculated for each specimen. The

FOREWING LENGTH: TAIL LENGTH RATIO

JoURNAL OF THE LEPIDOPTERISTS SOCIETY

SPRING SUMMER:

Fe O
Oo sues
O
~ fou fe)
iS :
(@)
\ Po O es O
O
le}
L “
| r @)
L e) é)
@)

60 86. “lea” “IZer As I60 180 200° ZaaeZzae
DAYS AFTER 1 JAN.
Relation of forewing length : tail length ratio to date of capture for 59 speci-

mare ellus from Mississippi, showing separation into spring and summer
TL = 2.05 and 137 days after January (17 May).

VoLUME 24, NUMBER 3 179

FOREWING LENGTH, MM

SPRING SUMMER

we
e¢ & @

PE Siamiice day ico (cz lod 0G oR NG

ANGLE BETWEEN OUTER AND LOWER MARGIN OF
FOREWING, DEG.

Fig. 3. Relation of forewing length to angle between outer and lower margin of
forewing for 67 specimens of G. marcellus from Mississippi, showing separation into
spring and summer subgroups.

range for the 59 specimens was 1.43 to 2.91. These results are plotted in
Fig. 2 against dates of capture.

Tail Pattern: Two types of tail pattern were recognized: (a) only the
tip of the tail white; (b) white extending far up each side of the tail
above.

Red Spot or Spots at Anal Angle of Hind Wing: Four variations of this
character were recognized : (a) red spot wide, covering the space be-
tween three veins, and step-shaped; (b) red spot of same dimensions as

180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

in (a) but divided by a narrow vertical black line along the intermediate
vein into two red areas each essentially square; (c) red spot reduced so
as to occupy the space between only two veins, and rounded; and (d)
red spot similar to (c) but accompanied by a much smaller satellite spot
in the adjacent space.

Submarginal White Lunules on Hind Wing: The number of prominent
white submarginal lunules was found to range from two to four. When
only two or three were prominent, often one or two were faintly indicated.

Fifth Black Stripe on Forewing: The middle, fifth, black stripe on the
forewing above extends downward only through the cell. Three forms of
this stripe were noted: (a) a band of generally uniform width, (b) a
band narrowing sharply away from the costal margin, and (c) the band
reduced to a spot.

Angle between Outer and Lower Margins of Forewing: The angle be-
tween the outer and lower (inner) margins of the forewing was measured
for each specimen. The measurements were made using a contact goniom-
eter.’ The relation between these results and forewing length is plotted
in Fig. 3.

Curvature of Outer Margin of Forewing: The outer margin of the
right forewing was inspected from above and classified as “concave” or

convex .

Discussion

Variation: A principal type of variation in G. marcellus is seasonal.
Brown (1965) quoted Boisduval’s letter of 1 June 1873 to W. H. Edwards
in which he wrote: “I believe you are right to put together Ajax and
Marcellus in spite of the difference there is in the coloration of the larvae
and the length of the tail of the insects in the perfect state [imago]. In
spite of the opposite opinion of Abbot, they are probably only seasonal
variations. The data on the Mississippi sample were therefore arranged
chronologically by date of capture and were examined together with the
relationships plotted in Fig. 1, 2, and 3 to determine if seasonal subgroups
could be segregated, and if so, how many such subgroups appeared to be
clearly differentiable and where the subgroup boundary or boundaries

should be drawn both with regard to characters and time. This examina-
tion led to the conclusion that the population represented by this sample
lent itself better to the drawing of one rather than more than one boundary

and hence was more clearly separable into two rather than to more than
(wo seasonal forms. The boundary that appeared most reasonable to draw
on the basis of change in characters corresponds, on the time scale, to the

lodel B, designed by S. L. Penfield, patented 31 July 1900, New Haven, Conn.

VOLUME 24, NUMBER 3 WSL

date of 17 May; one of three specimens taken on that date being assign-
able by characters to the earlier “spring” group; the other two being as-
signable to the later “summer” group. The final versions of Fig. 1, 2, and
3 were then modified using different symbols for specimens assigned to
the two groups. In each case a line can be drawn between areas in which
all the points representing specimens belonging to the two groups fall.

The two seasonal forms as segregated from the present sample may be
characterized and compared as follows:

Earlier “Spring” Later “Summer”

Dates taken 9 Feb-17 May 17 May-27 Aug
Forewing length, mm 29-39 36-47
(FWL)
Tail length, mm (TL) 11-19 18-29
FWL/TL LVS 1.43-2.00
Tail pattern White at tip only White extending up sides
(5 exceptions )
Red spots on hind wing Broad Narrow (1 exception )
HW submarginal lunules 3 + to 4 Five have fewer than 3 +

FW margin angle, deg. 102 or less (5 102 or more (3 exceptions )
exceptions )
Number of specimens So (25. &6, 80 (28 66, 7 22)
1299)

From the forewing characterization and comparison it is concluded that
the most unambiguously diagnostic characteristic is forewing length : tail
length ratio. Since the tail length increases with time more rapidly than
does the forewing length and since absolute dimensions are influenced at
any point in time by non-dependent factors such as sex and food, it is not
surprising that this ratio appears best to indicate the time-dependent
change that is observed. If this ratio is 2.00 or less the specimen belongs
to the later form regardless of its absolute size or other features. The
second most unambiguous character is the width of the red spot or spots
in the anal angle of the hind wing. If this area is broad, occupying the
spaces between three veins, the specimen belongs to the earlier form with
only rare exceptions (one in this series). All specimens having white on
the tail confined to the tip belong to the earlier form but, in this series,
five having white extending up the sides of the tail are also assigned to this
group. Since the sample includes comparable numbers of individuals
representing the two forms and since it includes individuals taken over a
period of 17 years, I believe it is as representative of the population
sampled as one of this size could be.

182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The nature of the curvature of the outer margin of the forewing does not
appear to be time dependent. It appears rather to be associated with the
sex of the specimen, as might be expected by analogy with other species,
but there are a significant number of exceptions to the association as is
shown in the following tabulation:

Curvature of outer margin Number of specimens

of right forewing, viewed from above 36 2&
concave 44 8
convex + 11

Variation in the shape of the fifth black stripe on the forewing above,
from a band of nearly uniform width to a band that narrows sharply down-
ward, appears to be random and not time dependent. The more extreme
reduction of this band to a spot was observed in four specimens, all belong-
ing to the later form (3 4 6,1 2) and, ina manner similar to the tendency
to obsolescence of some of the submarginal lunules on the hind wing
above, appears to be a character that is more likely to occur among in-
dividuals of that form but cannot be described as diagnostic of that form.

Nomenclature: Having discussed the results of the study of variation
within the sample of the Mississippi population it now seems appropriate
to discuss the names that have been used and the variation that has been
reported for populations of G. marcellus both throughout its range and in
specific portions of its range. Dos Passos (1964) gave the nomenclature as
follows:

“Graphium
269. marcellus (Cramer), “1779” [1777] (gen. vern. )

* ajax (Linnaeus), 1758 (partim) (Opinion 286)
walshii (Edwards), 1872
carolinianus (Holland ), 1931 (Opinion 259 )

form telamonides (Felder & Felder), “1864-67” [1864]

form floridensis ( Holland ), 1898
gen. aest. lecontei (Rothschild & Jordan), 1906”

and then records the names given to four aberrations.

Holland (1931, Plate XLIV) figured four specimens: 1, spring form

carolinianus Kdwards, = marcellus Cramer; 2. spring form floridensis
Holland (type); 3. summer form telamonides Felders, = lecontei Roth-
schild & Jordan; and 4. spring form walshii Edwards (type). The form

represented as # 4 “walshii” by Holland is similar to that which appears
carlier in Mississippi; the form represented as # 2 “floridensis” by Holland
which is quite similar to # 4, is matched by a few Mississippi specimens

VOLUME 24, NUMBER 3 183

but is not differentiable as a seasonal form in the present sample. The
form represented as # 1 “carolinianus” by Holland is typical of a very few
intermediate specimens from Mississippi that are rather inconsistent in
their characters. The form figured as # 3 “telamonides” by Holland is
similar to the later, “summer form as characterized above for the Mis-
sissippi series.

Most previous workers have discussed seasonal forms of G. marcellus in
terms of three such forms: smallest = earliest, intermediate both in time
and characters, and largest = latest. Clark (1932) wrote of the population of
the District of Columbia and vicinity as forms “marcellus,” “telamonides,”
and “lecontei’ having forewing lengths of 33-37 mm, usually 35 mm for
“marcellus,’ and 39-48 mm for “lecontei.” Field (1940) used the same
form names for Kansas and stated that the forewing of “marcellus” ranged
from 32-36 mm, averaged 34 mm, and the tails were 15 mm or less; “tela-
monides” had forewings averaging 37 mm; and “lecontei” (in Douglas
Co.) averaged 44 mm forewings and 23 mm or longer tails. Clark & Clark
(1951) wrote of the Virginia population that early spring individuals had
forewings usually 32-35 mm but up to 40 mm on the outer Coastal Plain.
They did not use form names for the earliest and latest forms but referred
to the late-spring form as “telamonides.” They also discussed the finding
of some individuals that closely resembled “the southeastern spring form
floridensis.” They stated that the summer form in Virginia had forewings
about 45 mm long. In no case has a report been found indicating the size
of the sample upon which the observations as to average or range in size
was based. All writers agree that as the season advances the butterflies get
larger and their tails get longer. No reference was found to the use of
forewing length : tail length ratio as a parameter. Kimball (1965) wrote
that the Florida population was “abundant throughout the state from
March to December. Morgan wrote *. . . of the three subspecies (sic) de-
scribed as differing slightly in size, hairiness, color, pattern, and length
of tails, and supposed to be restricted to certain seasons or regions, all may
be matched by Hillsborough County specimens throughout the year.
Fuller states that around Cassadega the species occurs in the three forms
in their usually recognized sequence; marcellus, February; walshi (sic)
(Edwards), March, and lecontei (R. & J.), June and July.” Kimball con-
cluded: “.... the whole subject of subspeciation in marcellus needs to be
worked out.” Forbes (1960), referring to New York and neighboring
states, mentioned an “early spring form (marcellus )” and a “summer brood
(lecontei)” adding “intermediate late spring specimens are telamonides
Felder” having some characteristics like the earlier and some like the later
forms. Haydon (1933) stated that, in Maryland, there were “three dis-

184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tinct seasonal forms”: marcellus 2.50-2.75 in., telamonides 2.75-3 in., and
lecontei 3-3.25 in. Weed (1926), likewise referred to “three distinct
forms”: marcellus, small early spring with short tails that show white only
on the tips; telamonides, late spring, larger, tails a little longer, and with
more white on outer half; and ajax, summer, larger, with very long tails.
Comstock and Comstock (1940) wrote of “three distinct forms,’ early
spring (marcellus) 2.6-2.8 in., tails 0.6 in., tipped with white = walshii
Edwards; late spring (telamonides), a little larger, tails % longer, bor-
dered by white on distal % to % of length; summer (ajax) 3.2-3.5 in., tails
7 longer than early spring.

Although, as has been noted above, most previous workers have dis-
cussed seasonal variation in terms of three “distinct” forms, no illustration
designated as representing the intermediate form was found in the litera-
ture. The intermediate form has usually been called “telamonides” but the
only figure designated “telamonides” that I found was that given by Hol-
land (1931) who refers to it as “summer form = lecontei.” A total of
eleven figures were found. The references to these figures, the measured
forewing and tail lengths and the computed forewing length : tail length
ratio are tabulated below:

FWL, Tbs FWL/ NO.
FIG.
Reference mm mm ATL 4
Holland (1931), plate XLIV, fig. 1 2
spring 38 16 2.38 I
fig.2 ¢ spring “‘floridensis” 38 HES 2.53 2
fig. 3 ¢ summer “telamonides” 44 24 1.83 3
fig. 4 @ spring “walshi” 34 14 2.43 4
Comstock & Comstock (1904), plate
V, fig. 1 spring oil 14 2.64 5
fig. 2 summer Al 23 1.78 6
Klots (1951), plate 24, fig. 7 ¢ spring
| (Pennsylvania ) PALL (B35) )) 9(15) 2a i
fig. 8 $ summer ( Alabama) “lecontei” 26( 43) 17(28) 1.54 8
Howe (1964), plate 9, fig. 4 (summer) 46 24 1.91 g
Clark (1932), plate 48, figs. 1, 2 9
summer ( Maryland ) 45 25 1.80 10
plate 49, figs. 1,2 2 spring
(Maryland ) 33 14 2.36 a
‘ Values in parentheses are computed with allowance for stated reduction in published size of

firure.

The forewing lengths and tail lengths tabulated above are plotted on Fig.
| which is drawn to the same scale as Fig. 1 and on which are shown
‘lines of the limits of the data for the Mississippi sample. There is

or considering the establishment of an intermediate group from

VOLUME 24, NUMBER 3 185

Boa ° O34 "telarionides
% )

O8!

!
?
)
y)

ne pridonsis 2 ALAN :
AS /
oe /

a a

7

“i
A7 PENNSYLVANIA
Aad oo
if
A \i MARYLAND

FOREWING LENGTH, MM

SPRING SUMMER
A oO

vor. 12 |4- 1G (Qo ey EOS EY KE SNS)
TAIL LENGTH, MM

Fig. 4. Relation of forewing length to tail length for 11 specimens of G. marcellus
figured in the literature and showing the boundaries for these parameters of the availa-
ble data on the Mississippi population taken from Fig. 1.

these data than from the Mississippi data. The range in FWL/TL ratios
for the eleven figured specimens and for the Mississippi sample are com-
pared below:

Spring Summer
Figured specimens 1.54-1.91 2.33-2.64
Mississippi sample 1.43-2.00 2.05-2.91

Distribution: Ehrlich & Ehrlich [1961] gave the range of G. marcellus
as “Florida to Texas, northward to Canada (rare in northern part of
range).” Macy & Shepard (1941) gave it as “from southern New England
through most of the United States east of the Rocky Mountains. . . . re-
ported several times from southern Michigan north to Saginaw Bay...”
Macy (in Macy and Shepard, 1941) reported seeing one in Hennepin Co.,

186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Minn. on 3 July 1932 and recorded Franzen’s report of one having been
taken in downtown Minneapolis. Sorenson (1967) states that only a single
specimen is known from Minnesota, from Mankato, Brown Co. It was not
listed for Colorado by Brown, Eff, and Rotger (1957). Forbes (1960)
gave the range as “Conn. to IIl., and s to Fla.” Fiske (1901) stated that
it was “said to have been taken in Hooksett (New Hampshire) but I
have nothing more than mere hearsay to go by.” Shapiro (1966) reported
taking 46 in the Delaware valley in 10 years. Klots (1951) gave the range
as southern New England (rare) west through southern Ontario, Mich.,
Minn., and Wisc., south through Central Florida and Gulf States. Field
(1940) wrote: “Canada south to Florida and west through the eastern
half of Kansas, Oklahoma, and Texas.” He gave records from eight Kan-
sas counties. Riotte (1967) noted that a specimen had been taken at
Humber Plains west of Toronto in 1891 but not since. Masters (1967)
listed it as abundant to common in northeastern Arkansas. Kendall (1964)
reported it as found from late March to early July in east Texas. He col-
lected eggs, larvae, and foodplant (Asimina parviflora (Michaux.) in
Polk Co., Texas, on 14 April 1962, and reared nine adults that emerged on
18-19 May, 6 June, 31 Aug 1962, and 25, 27 March and 11 April 1963. It
was not listed for the Waco, Texas, area by Gooch & Strecker (1924).
Lambremont (1954) and Ross & Lambremont (1963) recorded its occur-
rence in seven Louisiana Parishes; Kendall (1964a) added an eighth; the
flight period in Louisiana, based on these reports, is 17 March to 10 Sep-
tember. Harris (1950) reported it abundant over the entire state of
Georgia, February to October.

After reviewing these somewhat differing statements, I have concluded
that the available data indicates that G. marcellus occurs throughout
Florida (Kimball, 1965) rather than only to central Florida (Klots, 1951)
and generally in an area bounded by the Atlantic Coast northward at
least to Connecticut (Forbes, 1960) and westward, possibly into New
Hampshire (Fiske, 1901), through Toronto, Ontario (Riotte, 1967),
Michigan (Macy & Shepard, 1941), and Minnesota (Sorenson, 1967);
southward to include the eastern halves of Kansas and Oklahoma (Field,
1940 ) and east Texas (Field, 1940, Kendall, 1964).

Summary

A sample of the Graphium marcellus (Cramer) population of Missis-
sippi, consisting of 67 specimens (48 2 4, 19 2 2 ) taken on dates between
9 February and 27 August between 1951 and 1966 at localities in 12 coun-
listributed over the state, was examined. Data were obtained on

ig length, tail length, forewing length : tail length ratio, distribution

VOLUME 24, NUMBER 3 187

LEGEND

i

NUMBER

SPRING

FORE
WING

1s 2o 2s Bo 35 40
LENGTH, MM

Fig 5. Distribution of forewing lengths and tail lengths of Mississippi G. marcellus,
divided into spring and summer subgroups.

of white at tail tip, variation of red spot or spots at the anal angle of hind
wing, angle between outer and lower margin of forewing, curvature of
outer margin of forewing, variation in submarginal lunules of hind wing,
and variation of form of fifth black stripe of forewing. It was concluded
that the major variation is time dependent, that the sample may most logi-
cally be considered as including representatives of two (rather than more
than two) seasonal forms, that the most unambiguous character for sep-
arating these forms is forewing length: tail length ratio; that the change
from the earlier form to the later form occurs when this ratio drops to 2.00
or less which corresponds in time to approximately 17 May. Concurrently,
but less unambiguously, the forewing length increases from 37 mm or less
to 38 mm or more, the tail length from 18 mm or less to 19 mm or more, the
white at the tail tip extends up the sides of the tail, the red spot at the
anal angle of the hind wing narrows from being two spaces wide to being
one space wide, and the angle betwen the outer and lower margin of the
forewing increases from less than 102° to more than 102°. These two
forms may be referred to as earlier and later, or “spring” and “summer,” or

188 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

as gen. vern. “marcellus” and gen. aest. “lecontei.” A character that may
conveniently be used in sexing specimens is described. Conflicting state-
ments about geographic range are reviewed. It is concluded that G.
marcellus occurs throughout Florida and generally northward at least into
Connecticut, possibly into New Hampshire, westward to Toronto, Ontario,
Michigan, and Minnesota; southward to include the eastern halves of
Kansas and Oklahoma, and into eastern Texas.

Literature Cited

Brown, F. M., 1965. Three letters from J. A. B. D. de Boisduval to W. H. Edwards
and the identity of Melitaea pola Bdv. and Melitaea callina Bdv. Jour. Lepid. Soc.
19:197-211.

Brown, F. M., D. Err, AND B. Rorcrer, 1957. Colorado butterflies. Denver Mus.
Nat. Hist. Proc. 3—7 368 pp.

Crark, A. H., 1932. The butterflies of the District of Columbia and vicinity. U.S.
Nat. Mus. Bull. 157. 337 pp.

Crark, A. H., 1936. The swallowtail butterflies. Smithson. Ann. Rept. for 1935.
Publ. 3367:383—408.

Criark, A. H. Anp L. F. Ciarx, 1951. The butterflies of Virginia. Smithson. Misc.
Coll. 116(7), 239 pp.

Comstock, J. H. anv A. B. Comstock, 1904. How to know the butterflies. Appleton
& Co., New York. 311 pp.

Enrcuicu, P. R. anp A. H. Enruicu, [1961]. How to know the butterflies. Wm. C.
Brown Co., Dubuque, Iowa. 262 pp.

Firtp, W. D., 1940. A manual of the butterflies and skippers of Kansas (Lepidoptera,
Rhopalocera). Bull. Univ. Kans. 39. 328 pp.

Fiske, W. F., 1901. An annotated catalogue of the butterflies of New Hampshire.
N. H. College, Ag. Exp. Sta. Tech. Bull. 1. 80 pp.

Forses, W. T. M., 1960. Lepidoptera of New York and neighboring states, Part IV,
Agaristidae through Nymphalidae, including butterflies. Cornell Univ. Ag. Exp.
Sta. Mem. 371. 188 pp.

Goocn, W. T. ANp J. K. Srrecker, 1924. A list of diurnal Lepidoptera from the
vicinity of Waco, Texas. Baylor Bull. 27:21-28.

Harris, L., Jr., 1950. The butterflies of Georgia. Ga. Soc. Naturalists Bull. 5. 33 pp.

pe F.S., 1933. The Papilionidae of Maryland. Proc. Nat. Hist. Soc. Md. Vol.
2,14 pp.

HotLtAnp, W. H., 1931. The butterfly book (revised edition). Doubleday & Co.,
Garden City, N. Y. 424 pp.

ewe W. H., 1964. Our butterflies and moths. True Color, N. Kansas City, Mo.

)8 pp.

KenpALL, R. O., 1964. Larval food plants for twenty-six species of Rhopalocera
(Papilionoidea ) from Texas. Jour. Lepid. Soc. 18:129-157.

KENDALL, R. O., 1964a. New distribution records for three species from Arkansas,

_ Louisiana, and Texas ( Hesperiidae, Papilionidae). Jour. Lepid. Soc. 18:190-191.

KIMBALL, C. P., 1965. Lepidoptera of Florida. Div. of Plant Industry, Fla. Dept. of

Ag. Gainesville. 363 pp.

Kiors, A. B., 1951. A field guide to the butterflies. Houghton Mifflin Co., Boston.
349 pp.
LAMBREMONT, EH. N., 1954. The butterflies and skippers of Louisiana. Tulane Univ.

Stud. Zool. 1:127-164.
4. W., awd H. H. Sueparp, 1941. Butterflies. Univ. of Minn. Press Minne-

JU.

"
1A

VOLUME 24, NUMBER 3 189

Masters, J. H., 1967. Observations on Arkansas Rhopalocera and a list of species oc-
curring in northeastern Arkansas. Jour. Lepid. Soc. 21:206—209, 277.

MatTuHER, B. AND K. MatuHer, 1958. The butterflies of Mississippi. Tulane Univ.
Stud. Zool. 6:63—-109.

pos Passos, C. F., 1964. A synonymic list of nearctic Rhopalocera. Mem. Lepid.
Soc., No. 1. 145 pp.

Riorte, J. C. E., 1967. New and corrected butterfly records for Ontario and for
Canada. Jour. Lepid. Soc. 21:135-137.

Ross, G. N. anp E. N. LAMBREMONT, 1963. An annotated supplement to the state
list of Louisiana butterflies and skippers. Jour. Lepid. Soc. 17:148—158.

SHAPIRO, A. M., 1966. Butterflies of the Delaware Valley. Spec. Publ. Amer. Ent.
Soc. 79 pp.

SORENSON, J. T., 1967. The Rhopalocera of Minnesota, Part III, family Papilionidae.
Newsletter Assn. Minn. Ent. 1:56—57.

WEED, C.M., 1926. Butterflies. Doubleday, Page & Co., New York. 286 pp.

THE AEGERIID RAMOSIA FRAGARIAE IN A FLIGHT
TRAP, AND THE INTERPRETATION THEREOF

In mid-August 1966, my wife-and I were camped at Poker Flat, an attractive small
meadow at 5040 feet altitude in the forested Siskiyou Mountains of the northwest
corner of Siskiyou County, California. On the thirteenth I foolishly kept my eye on a
butterfly instead of on the ground, and ran over a small rock cliff. Fortunately I netted
the brute, Parnassius phoebus sternitzkyi McDunnough, on the way down, but in land-
ing head first on the talus lost considerable skin from arms, legs and side, and sprained
an ankle. Next day I was glad just to sit around camp.

Luckily a flight trap, the simple, one-pole, P. H. Arnaud-adaptation of a Malaise
trap, had been put up earlier. It was set on dry, rocky ground between the forest and
the marshy southern edge of the meadow, directly above the headwaters of the West
Branch of Indian Creek. To be doing something, I hobbled out to empty it every hour.
It caught quantities of flies and wasps throughout the day, but at the 3 PM servicing,
it contained also a series of a small black and orange clearwing moth, Ramosia fragariae
(Hy. Edwards). None was caught before or after the 2 to 3 PM period.

I interpreted this as a surprisingly restricted flight period for the species, and later
so reported it to several lepidopterist friends (I am a coleopterist ). But when the speci-
mens were readied for pinning and spreading a couple of years later, it was seen that
there were twelve males and one female, so it is more likely that the female inadver-
tently flew into the trap and was followed by her hopeful suitors.

This swarming of a number of males around one female may be characteristic of
these small clearwings. In my field notes for 26 July, 1964, referring to a spot near
Route 14, altitude 8,825 feet, 1.5 miles northeast of the summit of Granite Pass in the
Big Horn mountains of north central Wyoming, I recorded: “Near camp found a spot
where ¢ 6 aegeriids were swarming, wasp-like, over a tuft of grass; took a series, then
found a live @ in [the tuft of] grass; ants were attacking the 9. Got a couple more
@ 2 nearby.” These moths proved to be Ramosia chrysidipennis (Boisduval).

I am indebted to J. N. Shepard for identifying the butterfly, and to J. W. Tilden for
the names of the moths.

Hucu B. Leecn, California Academy of Sciences, San Francisco.

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NEW ASPECTS ON THE ASSOCIATION BETWEEN LYCAENID
LARVAE (LYCAENIDAE) AND ANTS (FORMICIDAE,
HYMENOPTERA)

Hans MALIcKy

Limnologisches Institut der Osterreichischen Akademie der Wissenschaften,
A-3293 Lunz, Austria

Introduction

The association between lycaenid larvae and ants is well known and has
been discussed several times in the literature. The most recent survey is
by Hinton (1951). Since the majority of published reports are purely
descriptive it would seem both desirable and necessary to consider the
evolutionary sources of the association using an experimental approach.
It is the purpose of this paper to present a summary of the essential facts
and new thoughts bearing on this association based on this approach; de-
tails of these investigations will be published elsewhere.

Current Hypotheses

The best known hypothesis advanced to explain the relationship be-
tween ants and the larvae originated with Thomann (1901). He suggested
that ants are able to drive away the caterpillar’s parasites, so that the
lycaenids benefit from the presence of the ant. In order to attract the ants
and keep them in attendance, the caterpillars exude a honey-like liquid
from a particular glandular organ, which was first described by Newcomer
(1912). According to this hypothesis the association is a symbiosis. This
hypothesis, however, has several weak points. Firstly, it has not been
confirmed that ants are able to drive away parasitic Ichneumonidae and
Tachinidae or even notice their presence. Secondly, there is no evidence
that the presence of the ant lowers significantly the rate of parasitism of the
caterpillars. Field observations by this writer show that the degree of
parasitism of lycaenid larvae fluctuates widely, as is usual in Lepidoptera,
whether ants are present or absent. Thirdly, the behavior of ants in respect
to lycaenid larvae shows no significant difference between species which
lack Newcomer's gland and those which have it.

\ second hypothesis has been advanced by Lenz (1917). According to
iim, the larval adaptations function to prevent aggression by the ant to-
vard the lycaenid larvae, and the honey-like liquid is exuded as a “bribe”

ecent knowledge supports this hypothesis.

VOLUME 24, NUMBER 3 191

Present Investigations
Methods

According to Thomann’s hypothesis the association of ant and lycaenid
larvae should lower the rate of parasitism of the larvae in comparison with
larvae of the same kinds which are not attended by ants. The examination
of this supposition seems to be technically impossible. In field experiments
it is not possible to exclude only the ants without changing other factors.
Results of experiments under laboratory conditions, on the other hand, do
not take into consideration other important field circumstances. It would
seem necessary to divide complicated ecological phenomena as far as
possible into factors, and to determine whether the presumed connections
actually exist.

The behavior of more than 50 ant species in relation to about 35 species
of lycaenid larvae has been investigated under both laboratory and field
conditions. The anatomy of the larvae has been studied microscopically
in the usual manner using serial sections, which were stained with paf-
halmi. More than 60 species of lycaenid larvae and pupae, mainly Euro-
pean, have been dissected and/or investigated morphologically. Additional
ecological investigations have been carried out in the field in order to find
out the biotopes and strata inhabited by ants and caterpillars.

The particular behavior of ants and its releasing mechanism

When not excited, ants perform a slow antennal motion which may be
called “groping” (Tasten). At excitement, under certain circumstances,
their antennal motion becomes much faster and the antennae are bent into
an acute angle. This motion may be called “palpation” (Trillern). The
antennal positions of both motions are demonstrated in Fig. 1. Ants which
are visiting a lycaenid larva perform intensive antennal palpation. This
is symptomatic of the fact that the ants perceive the caterpillar as an object
distinct from surrounding stones, litter, etc. Therefore palpating may be
regarded as a sign for the particular association. The sucking of “honey”
which originates from Newcomer's gland, cannot be such a sign because
these glands are not present in many species which are attended in the
same particular manner by ants. Releasers of groping are a number of
common stimuli of less specific significance. Palpating may be released
in different ways, but in lycaenid larvae presumably only in chemical ones
because no particular mechanical structures could be detected on the
integument of the caterpillars where palpation takes place. The ants do
not palpate with the same intensity over the whole surface of the cater-
pillars. They prefer distinct spots whose situation differs according to the
species but is constant within it.

192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Ants (Lasius alienus Foerster) attending a lycaenid larva (Plebejus argyr-
ognomon Bgstr.). Notice the antennal positions: groping (extreme left, right above )
and palpating (central left).

In the majority of species examined, one of these spots for increased
rate of palpation is the dorsal part of the prothorax. Another is situated
laterally near the spiracles, principally those near the posterior part of the
body (esp. in Lycaeninae and Theclini). A third spot is on the dorsal
part of the 7th and 8th abdominal segments surrounding the mouth of
Newcomer's gland (esp. in many species of “blues” and Strymonini).
These spots contain accumulations of small epidermal glandular organs
which may be called “perforated cupolas” (Fig. 2). These organs are, like
Newcomer's organ, homologous to hairs. They consist of two cells, one of
which is homologous to the trichogen cell of the hair and is sometimes
interlaced by longitudinal cavities which enclose the enormous nucleus.
It must be supposed that they produce the volatile substance which re-
leases the ant’s palpation. The second cell, homologous to the tormogen
cell, forms a plasmatic tube which serves as pipe-line for the secretion.
The distal part of the organ consists of a cuticular sclerotized perforated
plate of about 20 ym in diameter.

The “perforated cupolas” and their secretion, which acts as pheromone,
are of major importance for the association between lycaenid larvae and
ants. With the single exception of the European Nemeobius lucina L.,
these organs are present in all lycaenid larvae investigated. In some of
them (e.g. Feniseca tarquinius, Apodemia mormo, Phasis thero, and
Poecilmitis thysbe) the perforation of the distal plate can not be rec-
ognized distinctly with the equipment used, so that investigation with an
electron microscope is necessary. For comparison, caterpillars of several
other families have been investigated, but nowhere could perforated

I at

‘ected. They are likewise present in the pupae of lycaenids
d by ants in the same manner as the larvae.

VOLUME 24, NUMBER 3 193

Fig 2. Section of the dorsal part of the 7th abdominal segment of a lycaenid larva,
with glandular organs. Center: Newcomer’s gland with four glandular bladders, exuda-
tion duct (ed) and retractor muscles (rm). Right: “perforated cupolas” section.
Above: external parts of several cupolas.

Lack of “beat reflex” in lycaenid larvae

Experiments indicate that when the majority of non-lycaenid lepidopter-
ous larvae are physically disturbed, they perform a reflex lateral beating
motion. The biological function of this motion is presumably to shake
away rapacious arthropods. Nevertheless, this has little efficiency against

194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

attacks of ants since fast motions usually release aggression in ants. In the
lycaenid larvae examined (with the exception of Nemeobius lucina and
Phasis thero) this reflective motion is not performed. This behavior is
an additional adaptation to the association with ants.

Interrelations between the construction of integument of the caterpillars
and the mandibles of ants

The lycaenid larvae examined have, without exception, soft but tough
cuticles of unusually thick dimensions. Caterpillars other than lycaenids
of the same size have cuticles about 5-10 ym, but in full grown lycaenid
larvae, 200-300 pm thicknesses are usual (Fig. 3). Except for a thin
epicuticular layer of less than 1 wm, the integument consists of endocuticle
only. The exocuticle is restricted to local sclerotizations, e.g. those in the
dorsal part of the prothorax, the head capsule, the thoracic legs, the hooks
of abdominal legs and the like.

In behavior experiments, it often happened that ants attacked the lycae-
nid caterpillars despite the presence of perforated cupolas and Newcomer's
glands but, as a rule, the caterpillars were not damaged because of the
particular construction of the ant’s mandibles. This mandible construction
is unique within predacious arthropods. The molar part, a ledge with a
row of small teeth, prevents the penetration of the sharp, elongate incisive
part through the thick cuticle. Thin cuticles of caterpillars other than
Lycaenidae may be easily penetrated. On the other hand, the construction
of the mandibles of predacious Carabidae, Staphylinidae, and Silphidae,
and the chelicera of spiders, penetrate even this voluminous lycaenid
cuticle without difficulty.

The thickness and plasticity of the cuticle is, therefore, a goal adaptation
specifically for ants since it does not protect from attacks of other pred-
ators. Ant mandibles without this molar ledge also exist (e.g. in Dory-
linae and in the social parasites Polyergus and Strongylognathus etc.),
but their owners are not associates of lycaenid larvae.

In addition to its thickness, the lycaenid cuticle is folded in a particular
manner, to which fact the larvae owe their woodlice-like appearance.
Very vulnerable organs, such as the dorsal blood vessel and ventral nerve
cord, are protected under concavities (see Fig. 3). The prominent protru-

sions contain less important structures such as fat bodies etc. A biting ant
is forced to crush one of the prominent cuticular ledges or protrusions be-
tween its mandibles, thus giving additional protection to the caterpillar,
because its important organs become folded inward and not damaged. It
is noteworthy that the size of these ledges corresponds well with the size

Pad fry hae ene Pay ese Als ae s
he ma dies of medium-sized ants but not with those of other cater-

s, which are much larger (Fig. 3).

VOLUME 24, NUMBER 3 195

Fig. 3. Cross-section of a lycaenid (above), a nymphalid (center) and a noctuid
(below) larva, each with comparative cuticular thickness. Lateral figures are outlines
of ant heads. Left from top to bottom: Myrmica scabrinodis, Cremastogaster scutel-
laris, Solenopsis fugax. Right: Leptothorax lichtensteini, Camponotus ligniperda
(small worker), ditto. (big worker), Prenolepis nitens. Above: Outline of the head
of a caterpillar-hunting beetle (Calosoma inquisitor, Carabidae). Drawn to the same
scale.

196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Presence of particular organs in lycaenid larvae (only new results,

literature records are not included).

Species

Spalginae
Feniseca tarquinius F.

Aphnaeinae
Phasis thero L.
Poecilmitis thysbe L.

Theclinae
Thecla betulae L., Quercusia quercus L.,
Calycopis beon Cram.
Strymonidia acaciae F., Fixsenia pruni L.,
Strymon melinus Hb.
Strymonidia spini Den. et Schiff., S. w-album
Knoch, S. ilicis Esp., Callophrys rubi L., C.
avis Chapm., C. henrici Grote et Rob., C.
gryneus Hb., C. nelsoni Bdv., “Thecla’
palegon Stoll
Tomares ballus F., T. callimachus Ev., T.
mauretanicus Lucas

Lycaeninae
Lycaena phlaeas L., L. thoe Guer.,
+ 7 additional European spp.

Zizeeriinae

Zizina labradus Godt.

Lycaenopsinae
Lycaenopsis argiolus L.

Everinae
Everes comyntas Godt., E. argiades Pall.
Cupido minimus Fuessly

Lampidinae
Cosmolyce boeticus L.

Glaucopsychinae
Scolitantides orion Pall., Philotes vicrama
Moore, Glaucopsyche alexis Poda
Tolana iolas O., Maculinea, 4 European spp.

Plebejinae
Plebejus icarioides Bdv. + 3 addit.
Muropean spp., Aricia 4 spp., Cyaniris
semiargus Rott., Polymmatus 9 spp.
Hamear inae

\ lemeobius lucina | ae

Perforated
cupolas

+

4+

++

++

+

eans absent, R means reduction stages.

Newcomer's
gland

R?

++

++

Tentacles

an

VOLUME 24, NUMBER 3 197

Fig. 4. Part of a glandular cell (“trichogen cell”) of Newcomer’s organ of a lycaenid
larva (Cupido minimus) with extremely divided nucleus. Stained with KES (aceto-
carmine ).

The role of Newcomer's gland

The larger glands whose external mouth is situated dorsally on the 7th
abdominal segment were mentioned by Guenée (1867) and described
anatomically and histologically by Newcomer (1912), Ehrhardt (1914),
and Fiori (1958). Although the descriptions of these authors do not agree
in detail, I found that the construction of the organ is the same in all
species which I have investigated (Fig. 2). Ehrhardt noticed that the vast
glandular bladders consisted of only two big cells each with one branched
nucleus (Fig. 4) and not, as Newcomer and Fiori presume, of an epithe-
lium of many small cells or a syncytium. Additional details, including the
ontogeny of these organs, are given by Malicky (1969).

Newcomer's organ is not present in all species of lycaenid larvae. Mem-
bers of the same subfamily, tribe or even genus may have or lack this
organ (Table 1). Some species, e.g. Strymon melinus Hb., Strymonidia
acaciae F. and Fixsenia pruni L. appear to be actually in a stage of reduc-
tion of the organ. It may be presumed that this organ was present in all
ancestral lycaenids (except for some “riodinids’) and has been sub-
sequently reduced or lost in all lycaenid caterpillars where it does not
occur now.

It is well known that attending ants imbibe the honey-like liquid which
is exuded by the glands. The organ and its secretion plays an important
role within the association. Nevertheless, the lack of Newcomer’s organ in

198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

many species suggests that the secretion of the perforated cupolas is more
effective in the relationship between the two kinds of insects.

The function of the tentacles

The paired eversible cylindric organs on the 8th abdominal segment of
some lycaenid larvae have been known for a long time. They may be
termed “tenacles.” D...S (1785) noted as early as the eighteenth century
the “rote Warzgen” in the larva of Plebejus argus. Several authors sug-
gested that these tentacles have an odoriferous function which might signal
the presence of a honey-producing caterpillar to ants. I can not confirm this
suggestion. I have never noticed any significant reaction of ants to the ten-
tacles, and I could not find any glandular structures in, on or near them.
They contain only distally inserted setae of usual construction but minor
size. Therefore, the function of the tentacles still must be regarded as
unknown. They may be rudimental structures, developing from organs
such as Ross (1964), Bourquin (1953) and Bruch (1926) found in several
Ancylurinae and Hamearinae.

Which kinds of ants may be associated with lycaenid larvae in the field

In my experimenis, only European representatives of Formicinae,
Dolichoderinae, Myrmicinae, and Ponerinae were studied. Only five of
them proved to be incapable of reacting specifically to lycaenid larvae.
They were: one extremely specialized social parasite ( Polyergus rufescens
Latr. ), three specialized grain collectors (Messor mutica Nyl., M. rufitarsis
F., Aphaenogaster subterranea Latr.) and two soil-inhabiting species
(Myrmecina graminicola Latr., Ponera coarctata Latr.). The majority of
species investigated, such as members of the genus Formica, Camponotus,
Lasius, Myrmica, Solenopsis, Leptothorax, etc. showed the same reaction
in qualitative respects. Field data furnished by Downey (1962) from the
Nearctic Plebejus icarioides (Bdv.) help confirm this, since no less than 11
different ant species of diverse taxonomic affinities were found attending
one species of butterfly. In general, the ant species which attend lycaenid
larvae seem to be the same ones which also attend Homoptera and which
have a high-developed trophallaxis between adults and larvae. This hypoth-
esis may contribute to an explanation of the evolution of the association
between ants and lycaenids.

Which kinds of lycaenid larvae are attended by ants in the field

\ | |
if? +}

ingle exception of Nemeobius lucina, all species investigated
‘ory induced an ant association. Several species were fre-
‘ensively by the ants but the behavior of the ants toward

VOLUME 24, NUMBER 3 199

the larvae was the same in a qualitative sense. This result does not agree
with field notes, because while several kinds of lycaenid larvae (e.g. Ple-
bejus argus L., Scolitantides orion Pall.) are never found without ants,
other species have never been recorded as being ant-attended. Therefore,
additional field research was undertaken which led to the idea that for the
association of certain partners, a common biotope preference is most im-
portant. There is an additional preference of activity of certain kinds of
ants and caterpillars for the soil surface (e.g. in Tetramorium caespitum
L., Lasius flavus F., and Plebejus argus L.), for the herbaceous vegeta-
tion (e.g. in Myrmica scabrinodis Nyl. and several Polyommatus and
Lycaena spp.) or for the shrub and tree stratum (e.g. in Colobopsis trun-
cata Spin., Dolichoderus quadripunctatus L., and several Theclinae ).

Exceptions

As already mentioned, the majority of lycaenid larvae and ants may be
associated only in a facultative way. The caterpillars do not need the ant’s
presence. Nevertheless, in a few cases, close relations exist between
specific lycaenids and ants. In Europe, the four species of the genus
Maculinea are associated with ant species of the genus Myrmica. These
caterpillars are phytophagous up to the last molt and are then carried into
the ant nests by worker ants. There they live as social parasites, feed on
the ant’s brood, and are fed orally by the adults. This association is based
upon additional adaptations. The molts take place at very short intervals
so that the last molt occurs only a few days after the larval emergence
from the egg. I presume that this is necessary for the accumulation of the
highest number of perforated cupolas at the time when the caterpillar is
ready to be carried into a Myrmica nest. It therefore resembles the ant
larva in size, consistence and presumably odor. In addition, the critical
stage to be carried into the nest is well synchronized with the activity of
the host: in July and August, in the biotopes of these lycaenids, only mem-
bers of the genus Myrmica are in full activity, because the majority of
other Central-European ants have very low activity during summer. An-
other adaptation is that the Maculinea larvae perform a distinct begging
behavior similar to that of the ant’s larvae, which releases the feeding
response in the worker ants.

In the literature several lycaenids are also recorded to have carnivorous
associations, e.g. Lachnocnema bibulus F. (Cripps and Jackson, 1940),
Niphanda fusca Brem.-Grey and Spindasis takanonis Mats. (Iwase, 1953,
1955), Euliphyra mirifica Holl. (Lamborn, 1914) and Liphyra brassolis
Westw. (Chapman, 1902). However, such association depending in an
obligate way on the ant’s care is exceptional. Bionomical data are availa-

200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ble for about 800 lycaenid species, but only 10 are known to belong to
these exceptions.

Principles, function and evolution of the association

The association between lycaenid larvae and ants is based upon several
ethological, morphological and chemical goal adaptations of the cater-
pillars to the behavior and the social biology of the ants. There is no
reference to any adaptation of ants to the lycaenid’s habits. Some of the
caterpillar’s adaptations are based upon copies of signals (Signalkopien )
which are important in the ant’s social life. The secretion of the perforated
cupolas is presumed to be similar to, or identical with, ant pheromones.
The Maculinea larvae copy a Myrmica larva in size and habits. The lack
of the beating-reflex and the presence of a thick cuticle are adaptations to
the aggressive behavior and to the mandible construction of ants. There-
fore, the biological function of the association is to protect the caterpillars
from the ant’s aggression. Protection from ant aggression is, however, not
restricted to lycaenids. Among insects and other arthropods many ways are
realized which serve the same purpose. Armoured cuticles (e.g. in many
Coleoptera ), dense pilosity (larvae of Arctiidae, Lymantriidae etc. ), chem-
ical repellents (Coccinellidae, Zygaenidae), construction of protecting
cases (Psychid and Tortricid larvae) and other devices prevent ant ag-
gression or render it ineffectual. In this respect, the particular association
of the lycaenids is only one of several realized chances.

The evolution of this association can only be sketched in broad outlines,
since the details of anatomy and histology are available only in few kinds
of caterpillars. Presumably the present day larvae originate from a kind
which differed from other lepidopterous larvae by its voluminous cuticle.
Larvae such as Nemeobius lucina may derive directly from this stage.
Another evolutionary pathway lead to the development of big and ever-
sible glandular equipment similar to Newcomer's organ, which are pre-
sumed to have been serial in their earliest ancestral stage. On this evolu-
tionary line the perforated cupolas developed which proved to be more
effective than the big glands, which were subsequently modified and
reduced. The result of this evolution is a caterpillar which is provided only
with perforated cupolas, as it is demonstrated by species of Lycaena and
several Theclini.

The well known carnivorous habits of many lycaenid larvae (Dethier
‘927, Clausen 1962) have no connection with the above mentioned evolu-
lion. It is certain that the obligate rapacious lycaenids evolved indepen-
dently from each other, originating their carnivorous habits from different
levels of ant adaptation. I agree with Brues’ (1936) ideas, but I can not
he Lamarckian suggestions of Balduf (1938).

VOLUME 24, NUMBER 3 201

Suggestions for future work

It is highly desirable that we investigate the anatomy and histology of
many additional kinds of lycaenid caterpillars, particularly additional sub-
families which are well represented in the Ethiopian, Indoaustralian and
Neotropic regions. It would be very revealing to research the details of
the specialized organs by means of ultra-optical methods. The chemistry
of the secretions and their interrelations with ant’s pheromones are still
unknown and need to be investigated. The behavior of additional ant
species, particularly those of other than European origin, may shed new
light on these relationships.

Acknowledgments

I am grateful to many colleagues who supported my work by contribut-
ing lycaenid material and in many other ways, above all to Dr. B. Antal-
falvi, Budapest; Mr. C. G. C. Dickson, Cape Town; Prof. J. C. Downey,
Cedar Falls (Iowa); Mr. N. W. Elfferich, Rotterdam; and Dr. H. Zwolfer,
Delémont (Switzerland). Many thanks are also due to Prof. Downey for
correcting and proof-reading the manuscript.

Summary

Comparative anatomical, histological and ethological studies in about
60 kinds of lycaenid larvae and pupae and ethological studies in about
50 kinds of ants were undertaken. The particular adaptations of the lycae-
nid larvae correspond with details of behavior, eidonomy and social
biology of ants and are regarded as protection from the ants’ aggression.
Individual species of lycaenids are dependent social parasites of ants, but
the majority are associated with ants only in a facultative way. True sym-
biosis is unknown. The evolutional pathway is suggested.

Literature Cited

Batpur, W. V. 1938. The rise of entomophagy among Lepidoptera. Amer. Nat.
72: 358-379.

Bourguin, F. 1953. Notas sobre la metamorfosis de Hamearis susanae Orfila, 1953
con oruga mirmecéfila (Lepid. Riodinidae). Rev. Soc. Ent. Argent. 16: 83-87.

Brucu, C. 1926. Orugas mirmecofilas de Hamearis epulus signatus-Stich. Rev. Soc.
Ent. Argent. 1: 1-9.

Bruers, C. T. 1936. Aberrant feeding behavior among insects and its bearing on the
development of specialized food habits. Quart. Rev. Biol. 11: 305-319.

CuapMan, T. A. 1902. On the larva of Liphyra brassolis Westw. Entomologist 35:
225-228, 2522.55.

CLAusEN, C. P. 1962. Entomophagous insects. New York: Hafner.

Cripps, C., Jackson, T. H. E. 1940. The life history of Lachnocnema bibulus (Fab. )
in Kenya (Lepid., Lycaenidae). Trans. Roy. Ent. Soc. Lond. 90: 449-452.

202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Deter, V. G. 1937. Cannibalism among lepidopterous larvae. Psyche 44: 110-
1S,

Downey, J.C. 1962. Myrmecophily in Plebejus (Icaricia) icarioides (Lepid.: Lycae-
nidae). Ent. News 73(3): 57-66.

D...S. 1785. Entomologische Fragmente. N. Mag. Liebh. Ent. 2: 383-384.

Enruarpt, R. 1914. Uber die Biologie und Histologie der myrmekophilen Organe
von Lycaena orion. Ber. Naturf. Ges. Freiburg, i. Br., 20: XC—XCVIII.

Front, G. 1958. “Strymon ilicis’ Esp. (Lepid. Lycaenidae). Boll. Ist. Ent. Univ.
Bologna 22(1957): 205-256.

GuENEE, M. 1867. D’un organe particulier que présente une chenille de Lycaena.
Ann. Soc. Ent. Fr. (4) 7: 665-668.

Hinton, H. E. 1951. Myrmecophilous Lycaenidae and other Lepidoptera—a sum-
mary. Proc. South Lond. Ent. Nat. Hist. Soc. (1949-50) pp. 111-175.

Iwase, T. 1953. Aberrant feeders among Japanese Lycaenid larvae. Lepid. News 7:
45—46.

1955. The sixth aberrant feeder in Japan—Spindasis takanonis (lLycae-
nidae). Lepid. News 9: 13-14.

Lamporn, W. A. 1914. On the relationship between certain West African insects,
especially ants, Lycaenidae and Homoptera. Trans. Ent. Soc. Lond. (1913) pp.
436—498.

Lenz, F. 1917. Der Erhaltungsgrund der Myrmekophilie. Z. indukt. Abst. Verer-
bungsl. 18: 44-46.

Mauicxy, H. 1969. Versuch einer Analyse der 6kologischen Beziehungen zwischen
Lycaeniden (Lepidoptera) und Formiciden (Hymenoptera). In preparation.
Newcomer, E. J. 1912. Some observations on the relations of ants and lycaenid
caterpillars, and a description of the relational organs of the latter. Jour. New

York Ent. Soc. 20: 31-36.

Ross, G. N. 1964. Life history studies on Mexican Butterflies. II. Early stages of
Anatole rossi, a new myrmecophilous metalmark. Jour. Res. Lepid. 3: 81-94.
THoMANN, H. 1901. Schmetterlinge und Ameisen. Beobachtungen iiber eine Sym-
biose zwischen Lycaena argus L. und Formica cinerea Mayr. Jb. Naturf. Ges.

Graubunden 44: 1-40.

OCCURRENCE OF COENONYMPHA INORNATA (SATYRIDAE) IN MAINE

It has been noted in recent years that Coenonympha inornata (Edwards) has
pice onpandiue its range in the northeastern portion of the United States and
sanada.,

On July 5, 1968 I collected a female of this species in a roadside bog between

Enfield and Passadumkeag, Penobscot Co., Maine. The specimen, which was taken
in the early afternoon, appeared to be about one to two days old. Poor weather
miu: and a subsequent collecting trip to Canada prevented further collecting
in this area.

The specimen has been determined by F. Martin Brown as being closest to C.

imornata inornata (Edwards). As far as can be determined, this is the first record
lor this species in the State of Maine.
FORD D. FrErris, Laramie, Wyoming.

VOLUME 24, NUMBER 3 203

DISTRIBUTIONAL NOTES ON THE GENUS MESTRA
(NYMPHALIDAE) IN NORTH AMERICA

JoHNn H. Masters?

P.O. Box 7511, Saint Paul, Minnesota

The genus Mestra Hiibner is of Neotropical distribution and contains six
species, one of which, amymone, ranges northward into the United States.
Mestra amymone (Menetries) is a breeding resident of the lower Rio
Grande Region of Texas, but has been frequently recorded as a “stray” far
north of here (figure 1).

Many tropical species, such as Phoebis philea (Johansson), are noted
for sometimes “straying” far northward, but the records for Mestra amy-
mone differ from all others in one important respect, the frequency of mul-
tiple captures in the extra-normal range. Records of tropical butterflies, in
regions considerably north of their normal range, consist almost exclusively
of single captures; yet nearly 50% of similar records for Mestra amymone
are for multiple captures. This is significant because it may indicate that
M. amymone is actually breeding in many of the areas where it has been
found.

The reported foodplant for Mestra amymone is a species of Tragia (fam-
ily Euphorbiaceae), a plant commonly known as “Noseburn” (because of
an effect the plants are alleged to have on livestock) or, in Mexico and
parts of the Southwest, as “Chichicastle.” Most workers have excluded any
possibility of M. amymone’s breeding north of its normal range on the as-
sumption that its foodplant, Tragia, is not available. Brown (1957) for ex-
ample, says: “There is no chance that this species will become naturalized
in Colorado. It is a tropical butterfly that breeds as far north as southern
Texas. Its foodplant, Tragia, does not grow in this region.” Johnson
(1962), however, refers to the genus Tragia as “. . . widespread in the arid
grasslands and brush of northern Mexico, north to Arizona, Colorado and
Kansas. ...” It is possible then that M. amymone can breed on Tragia far
north of southern Texas.

Noseburn is a small plant with small staminate flowers in racemes,
narrow serrate leaves and trailing vine-like stems covered with stinging
hair. Nine species and varieties of North American Tragia have been
described, but modern workers, such as Johnson (1962), McVaugh (1961 )
and Shinners (1961), concur that they all belong to a single variable
species, neptifolia Cavanilles. The approximate recorded range of Tragia

1 Research Associate, Section of Insects & Spiders, Carnegie Museum, Pittsburgh, Pa.

204 JouRNAL oF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Distribution of Mestra amymone (Menetries) in the Great Plains Region of
the United States. Solid circles indicate multiple captures (more than one specimen
in the same year); open circles, single captures. The cross hatched area indicates the
approximate permanent breeding area. The hashed line traces the approximate northern
limits of Tragia neptifolia Cavinilles, the larval foodplant for M. amymone.

neptifolia in the United States has been traced (hashed line) on the map
(fig. 1), it encircles most of Texas, Oklahoma and Kansas plus parts of Ari-
zona, New Mexico, Colorado, Nebraska, Iowa and Missouri—enclosing all
but three of the U.S. records for Mestra amymone. Within its range, Tragia
neptifolia occurs in almost all habitats from sea level to 8000 feet, but is
most abundant on rangeland at moderate elevations.

Until early stages are collected, there will be no proof that M. amymone
breeds north of Texas; in the meantime, the most convincing “circumstan-
tial” data that it does, comes from Reed (1913) who writes: “... in the
summer of 1907, I found quite a large colony of them [Mestra amymonel]
near Cordell, Washita County [Oklahoma]. I took specimens over a range
of about eight or ten miles in extent. They were permanently located and

VoLUME 24, NUMBER 3 205

were breeding.” Other records of multiple captures dot Oklahoma and
Kansas and there is a multiple record for South Dakota. We must assume
that M. amymone does indeed occasionally breed on the Great Plains, but
is unable to overwinter and become permanently established.

Mestra amymone is a weak flying species usually found in open areas,
and along roadsides and the edges of woods and is easily captured in flight
or at flowers. W. H. Howe (in litt.) has taken specimens in bait traps near
Ottawa, Kansas. Unlike most of the migratory butterflies it is not a strong
flier and in fact seems to be so fragile as to be unlikely to survive being
swept great distances by sheer force of the winds. I am of the opinion that
in some years overpopulation or drought may cause a shortage of food in
the normal range of the species, and the butterflies disperse northward
looking for foodplant. This theory would help explain the frequency of
records from widely separated areas in the same year, i.e. Colorado Springs
and Ottawa in 1950.

Records for Mestra amymone taken north of Texas may be summarized
as follows:

ARKANSAS: I took five specimens in less than an hour's collecting near
Texarkana, Miller County on August 31st, 1963.

COLORADO: Brown (1957) cited a specimen taken by Norman Mar-
ston at Hartman, Prowers County, one by himself in Bear Creek Canyon,
near Colorado Springs, July 4th, 1950, plus a possible site record near
Jimmy Camp Creek, southeast of Colorado Springs.

IOWA: The only Iowa specimen of M. amymone is an undated specimen
taken by Dr. L. G. Stempel near Macedonia and reported by Lindsay
(GISIEZ)). ,

KANSAS: Calkins (1932) recorded several specimens from Scott
County in August and September of 1931; Field (1938) reported records
from Sumner, Greenwood and Shawnee counties; Overman (1914) re-
ported a specimen taken during October 1904 near Lawrence, Douglas
County; Howe (1958) reported eight specimens taken at Ottawa, Franklin
County during 1950.

MINNESOTA: A single museum specimen has been located bearing
the data: “Pipestone, Minn. 1894.” This specimen may have originated
with Truman who made his collection at Volga, South Dakota (actually
not far away) at approximately this time. The present location of the
Truman collection is unknown.

NEBRASKA: The popular butterfly guides (i.e. Holland, 1930; Klots,
1951; and Ehrlich & Ehrlich, 1961) all record M. amymone as straying
north to Nebraska. However, Leussler (1938) excluded it from his state
list for Nebraska and there are no other published records from the state.
I have been unable to locate any specimens in museum collections. Ne-

206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

braska specimens will no doubt be collected at some date in the future; in
the meantime, it can be assumed that those specimens collected in Minne-
sota, Iowa and South Dakota either migrated through Nebraska or are
progeny of butterflies that did.

OKLAHOMA: Reed (1913) stated that M. amymone was a rare species
in Oklahoma, but could be found by the careful collector in almost any
year; his records include Washita (1907), Ponotoc (1905) and Kay (1911)
counties. The butterfly was also taken in moderate numbers in Comanche
County in 1950 and again in 1954.

SOUTH DAKOTA: Truman (1896) listed Mestra amymone among the
species that he had taken at Volga, South Dakota; undoubtedly these were
taken in a single year, probably 1894.

There are no known records of Mestra amymone for Arizona, New Mex-
ico or Missouri; Tragia neptifolia, the foodplant, occurs in these areas and
the butterfly may eventually be found as well. There is also a remote pos-
sibility that specimens of M. amymone will turn up eventually in Wyo-
ming or Louisiana. Distributional data for Texas and Central America fol-
low:

TEXAS: Gooch & Strecker (1924) recorded Mestra amymone as
double-brooded and very common at Waco, McLennan County, and this is
apparently the most northerly point where amymone has been known to
overwinter. Roy O. Kendall of San Antonio, Texas supplied me with a list
of all Texas localities at which M. amymone has been collected; these in-
clude Aransas Bandera, Bee, Bexar, Bastrop, Brewster, Cameron, Comal,
Crockett, Culberson, Dallas, Dimmit, Frio, Harris, Hidalgo, Jeff Davis,
Jim Wells, Kendall, Kerr, Kinney, Kleberg, Live Oak, McLennan,
Medina, Pecos, San Patricio, Starr, Tom Green, Swisher, Travis, Uvalde, Val
Verde, Victoria, Zapata and Zavala counties. As can be seen by the map
(fig. 1) these show a very strong concentration in the lower Rio Grande
Valley.

MEXICO: Hoffman (1940) cited the Mexican range of M. amymone as
temperate and hot areas over almost all of the Republic. Actually the
butterfly seems to be present throughout central and southern Mexico at

low or moderate elevations, but in northern Mexico is confined to that
area east of the Continental Divide.

CENTRAL AMERICA: The southern limit of Mestra amymone in
Central America is not known. It has been taken in Guatemala and British
Honduras by E. ©. VVelling and in El Salvador by M. Serrano. It probably
reaches Nicaragua an .sta Rica but, along with other members of the

genus, seems to be absent from Panama (Dyar, 1914; Huntington, 1932; GC.
Small, in litt.). In Sout! America, Mestra is represented M. bogotana

VOLUME 24, NUMBER 3 207

Felder and other species; the range “gap” in Panama does not seem to be
due to a lack of available foodplant.

Records of Mestra from Florida are undoubtedly erroneous despite the
fact that Strecker described Cystineura | Mestra] floridana from the Ever-
glades. According to Fox (1942), Strecker “deliberately misquoted the lo-
cality and number of specimens in order to conceal the fact that he had
used the Mengel series for his description”. Twelve specimens in the
Mengel Collection originated with Fred de Hart and bear the data label of
“Crystal River, Florida” [Citrus County?], a locality 70 miles north of
Tampa. Kimball (1965) removed M. floridana from the Florida list and
presumed either that they were not taken in Florida or that they may have
represented an isolated colony that was wiped out by the freeze of 1899.
This second theory is extremely doubtful because there is no foodplant,
Tragia, available in Florida. M. floridana is identical in appearance to
Mestra bogotana cana Erichson of Trinidad, and the most plausible ex-
planation is that the Mengel specimens originated there and were mis-
labelled. Mestra bogotana extends northward in the Antilles only to St.
Lucia and Dominica. A third Mestra, dorcas Fabricius, is found between
here and Florida on Jamaica. Archimestra teleboas Menetries, which was
formerly placed in Mestra, occurs in the Antilles also. Although Mestra
floridana still appears in our North American checklists (i.e. dosPassos,
1964), I am of the opinion that it should be “sunk” as a synonym of cana
and removed from future checklists.

Without getting in a discussion over the need for or the value of vernac-
ular names, it is interesting to mention that there is an old name, “The
Texas Bagvein”, that has not been used for Mestra amymone since the
1930’s. It is certainly a more imaginative name than “The Amymone”
which is currently in use in field guides.

I appreciate the assistance of Dr. George Wallace of the Carnegie Mu-
seum and Dr. Frederick Rindge of the American Museum of Natural His-
tory for allowing me access to their collections to obtain distributional
data, and of Roy O. Kendall of San Antonio, Texas for supplying me with
detailed distributional data for M. amymone in Texas.

Literature Cited

Brown, F. M., 1957. Colorado Butterflies. Denver Museum of Natural History. 368
pp.

Caxxins, V. F., 1932. The Rhopalocerous Lepidoptera of Scott County, Kansas. Ent.
News 43: 210-215, 225-229, 257-260.

posPassos, C. F., 1964. A synonymic list of the Nearctic Rhopalocera. Memoir no.
1, Lepidopterists’ Soc., 145 pp.

Dyar, H.G., 1914. Report on the Lepidoptera of the Smithsonian biological survey
of the Panama Canal Zone. Proc. U.S. National Mus. 47: 139-350.

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Eurucu, P. R. & A. H. Esruicu, 1961. How to know the butterflies. Wm. C.
Brown Co., Dubuque, Iowa. 262 pp.

Fretp, W. D., 1939. A manual of the butterflies and skippers of Kansas. Bull. U.
Kansas, 39(10), 328 pp.

Fox, R. M., 1942. Catalogue of the types in the L. W. Mengel butterfly collection.
Reading Public Museum, Sci. Pub. no. 4, 23 pp.

Goocu, W. T. & J. K. Srrecker, 1924. A list of diurnal lepidoptera from the vicinity
of Waco, Texas. Baylor Bull. 27(3): 21-28.

HorFMan, C. C., 1940. Catalogo sistematico y zoogeografico de los lepidopteros
Mexicanos. Primera parte. Papilionoidea. An. Inst. Biol. Mexico 11: 639-739.

Hotianp, W. J., 1930. The butterfly book (revised edition). Doubleday & Co.,
Garden City, New York. 424 pp.

Howe, W.H., 1958. Some new state Lepidoptera records for Kansas. Jour. Kansas
nts SOC} oles a iene

Huntincton, E.I., 1932. A list of the Rhopalocera of Barro Colorado Island, Canal
Zone, Panama. Bull. American Mus. Nat. Hist. 63: 191—230.

Jounson, M. C., 1962. The noseburn (Tragia, Euphorbiaceae) of western Texas.
Rhodora 64: 137-142.

KimsBa.1, C. P., 1965. Lepidoptera of Florida. Arthropods of Florida, Vol. 1, Florida
Dept. Agr., 363 pp.

Kuorts, A. B., 1951. Field guide to the butterflies. Houghton Mifflin Co., Boston,
349 pp.

Leussuter, R. A., 1938. An annotated list of the butterflies of Nebraska with the
description of a new species (Lepid.: Rhopalocera). Ent. News 49: 3-9, 76-80,
213-218, 275-280, 50: 34-39.

Linpsay, A. W., 1917. A list of the butterflies of Iowa. Ent. News 28: 347-353.

McVaueu, R., 1961. Euphorbiaceae novae Novo-Galicianae. Britonia 13: 145-205.

OverMAN, C.I., 1914. Cystineuwra amymone (Lepid.). Ent. News 24: 415.

Reep, E. L., 1913. Cystineura amymone (Lepid.). Ent. News 24: 279.

SHiINnNERS, L. H., 1961. Tragia nepetaefolia var. leptophylla instead of var. ramosa
(Euphorbiaceae). Southwest Nat. 6: 101.

TRUMAN, P. C., 1896. Lepidoptera of South Dakota. Ent. News 7: 298-299, 8: 27-—
29.

FREEMAN COLLECTION OF MEGATHYMIDAE DONATED TO
THE AMERICAN MUSEUM OF NATURAL HISTORY

lor the past 30 years Mr. H. A. Freeman has concentrated on the Megathymidae.
During this period he has worked out the life histories and food plants of most of the

species and subspecies in this family, and he has named many of these. His magnificent
collection, containing 2353 specimens, nearly all of them reared from the early stages,
has been donated to the American Museum of Natural History. It includes nine holo-
types, 14 allotypes, one neallotype, and 638 paratypes; 76 genitalic preparations and a
large number of pupal cases are also included. All of Freeman’s primary types of the
\iegathymidae, with one exception, are now in the collection of the American Museum
of Natural History. This generous donation gives the American Museum one of the
vugest and most complete collections of this family in existence.

erick TH. Rinnce, Curator, Department of Entomology, The American Museum

Vatural History, New York, New York 10024

VoLUME 24, NuMBER 3 209

THE EFFECT OF PHOTOPERIOD ON THE TERMINATION OF
PUPAL DIAPAUSE IN THE WILD SILKWORM, ACTIAS LUNA!

Davin A. WRIGHT
18 Clinton Place, Woodcliff Lake, New Jersey

Williams and Adkisson (1964) have demonstrated that photoperiod
controls the termination of pupal diapause in the silkworm, Antheraea
pernyi Guérin-Menéville. For example, at 25° C, short-day conditions (4-
to 12-hour photophases) strongly inhibit the termination of pupal dia-
pause, while long-day conditions (15- to 18-hour photophases) promote
the termination of diapause. The experiments which follow were carried
out to determine if photoperiod also controls the termination of pupal
diapause in the silkworm, Actias luna Linnaeus.

The first two experiments were carried out to determine the effect of
photoperiod on previously chilled pupae. Sixty A. luna pupae were pur-
chased from Butterfly Art Jewelry, Inc. on January 3, 1967 (I was told
that these were raised and stored outdoors). They were kept in the
garage where it was cool, until January 10 and then divided equally
among four 2 pound coffee cans. Each can was covered with mosquito
netting and immersed to a depth of about 2 inches in a fish tank of water.
A constant temperature heater kept the water at 26-28° C. The containers
were exposed to photophases of 0, 11, 16, and 24 hours. The 11-hour and
16-hour photophases were achieved by covering the containers with 3
pound coffee cans at 6 P.M. and 11 P.M. respectively and removing
these at 7 A.M. The 0- and 24-hour photophase containers were left
covered and uncovered respectively. Each of these latter cans was
painted black on the inside and placed so that air but no light could get
in. The whole set-up received illumination from two 40 watt GE F400W
fluorescent cool white lamps hung about 4 feet above the containers.

Pupae exposed to the 11-hour and 16-hour photophases emerged as
adult moths in an average of 17 days. The pupae exposed to continuous
darkness emerged in an average of 19 days and those exposed to con-
tinous light emerged in an average of 20 days (Table 1). The fact that
the first of these adults emerged in only 6 days as against 17 days in the
following experiment suggests that many were no longer in diapause at
the start of this experiment. If it can be assumed that those moths that
emerged on or after the seventeenth day were all in diapause at the start
of the experiment, the comparative times of emergence were such as to
suggest that photoperiod was not an important factor.

1 Part of a project which won the $8000 second prize in the Westinghouse Science Talent Search,
Washington, D.C., 1969.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. TOTAL ADULTS EMERGING FROM COMMERCIAL PUPAE®

0-hour 11-hour 16-hour 24-hour
Days Photophase Photophase Photophase Photophase

6 0 1 0 0
i 0 1 1 0
8 0 2 1 0
9 0 2 2 Z
11 Ph 2 Dp 3
12 ®) 3 2, 3
13 5 3 2 5
14 6 4 5 7
15) 7 5 7 8
16 8 7 8 8
iy 8 8 9 9
18 8 9 9 10
19 9 9 9 10
20 10 9 10 10
21 10 10 10 10
23 10 10 IL 10
24 I 10 ILL 10
26 11 10 2 10
ANS) 11 Il i 10
30 IAL 11 12 We
ol 2 11 2 12
34 13 IL 13 12
35 14 Wa Is} 12
39 14 Le ks) 13
40 14 Le 13 14

“The remaining specimens were dead.

TABLE 2. TOTAL ADULTS EMERGING FROM PUPAE IN DIAPAUSE*

11-hour
Photophase

0
2
3
)
6
7
8
8
9
10
10
10
10
12
12
13
14

ling specimens were dead.

16-hour
Photophase

OODAGAWIUIUIWNN WW

VOLUME 24, NUMBER 3 DAT

100

80

60

40

PERCENT EMERGED

20

0 i 16 24
PHOTOPHASE - HOURS

Fig. 1. The effect of photoperiod on the termination of diapause in the pupae of
Actias luna not previously chilled. Percentage emergence after: A, 8 weeks; B, 13
weeks; C, 18 weeks; D, 23 weeks; E, 28 weeks.

In the second experiment A. luna larvae were raised under an 11-hour
photophase as previously described by Wright (1967) to assure that all
pupae were in diapause. The cocoons were stored in the garage, still under
11 hours of illumination, from June until February 28 so that they would
be exposed to a period of low temperature before emergence. On Feb-
ruary 28, 15 pupae were put into each of two 2-pound coffee cans. The
procedure was essentially the same as that in the first experiment except
that a bath temperature of 24-25° C and photophases of 11 and 16 hours
were used.

The pupae exposed to the 16-hour photophase emerged as adult moths
in an average of 33 days and those exposed to the 11-hour photophase
emerged in an average of 29 days (Table 2).

A third experiment was carried out to test the effect of photoperiod on
unchilled pupae. Sixty diapausing pupae were bought from Bill Shibe of

212 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Moorestown, N. J. The cocoons were spun about September 2 and were
shipped 2 weeks later. The experiment was begun on September 21.
Fifteen A. luna each were exposed to 0-, 11-, 16-, and 24-hour photophases
at 24-25° C using the same apparatus described above. The pupae exposed
to the 16-hour photophase emerged in an average of 69 days and those
exposed to the 24-hour photophase emerged in an average of 152 days. At
the conclusion of the experiment (200 days), 67% of the pupae exposed to
the 11-hour photophase and 17% of those kept in darkness had emerged
(Claire, 1).

The difference in the number of days required for emergence of the
adult moths from previously chilled pupae under the various photoperiods
does not appear to be significant, and these experiments therefore suggest
that day-length has little or no effect on the termination of diapause of
overwintered A. luna pupae. On the other hand, the difference in the time
of emergence of the unchilled pupae is significant, and we may conclude
that photoperiod does play an important part in the termination of dia-
pause of such pupae.

These results are interesting because they suggest that in A. luna, photo-
period is important in the termination of diapause of unchilled pupa but
not of previously chilled pupae, whereas Williams and Adkisson demon-
strate that in A. pernyi photoperiod affects the termination of pupal dia-
pause of both chilled and unchilled pupae.

Literature Cited

WituiAMs, C. M., anp P. L. Apxisson, 1964. Physiology of insect diapause. XIV.
An endocrine mechanism for the photoperiodic control of pupal diapause in the
oak silkworm, Antheraea peryni. Biol. Bull. 127:511-525.

Wricur, D. A., 1967. The effects of photoperiod on the initiation of pupal diapause
in the wild silkworm, Actias luna. Jour. Lepid. Soc. 21:255-258.

FIELD NOTES ON THREE SKIPPERS IN TEXAS (HESPERIIDAE)

While collecting in the Rio Grande Valley-Brownsville area in southern Texas, the
author took three skippers rarely taken in the United States. All were collected in the
Santa Ana National Wildlife Refuge, Hidalgo County, Texas. The first, Synapte
salenus Mabille (9 ) was taken on the trail between the Old Spanish Cemetery and
the Inland Lake in a shaded, grassy, low spot on 27 August, 1968. Pyrgus albescens
Vloetz ( 4 ) was captured in a clearing at the river end of West Gate Road, 29 August,
L968. A female Pyrgus communis was also taken at the same time and place. On the
foo! trail between West Lake and West Gate Road, on 19 October, 1968, Celaenor-
minus stallingsi Yreeman was taken when it settled on a low plant growing in a patch

unshine coming through the trees overgrowing the trail. This is the second United

‘
Stote

. P ness , ray ee pe 5 ‘
‘tales record for this species. Determinations were made by H. A. Freeman, Garland,

josePH I. Doyie TI, 6127 Thunder, San Antonio, Texas 78238.

VOLUME 24, NUMBER 3 Zale

COMMENTS ON FORMS OF GONEPTERYX ASPASIA (PIERIDAE )
DESCRIBED BY SHU-ITI MURAYAMA

Yurrt P. NEKRUTENKO

Ukrainian Research Institute for Plant Protection, Vasilkovskaya Street 51, Kiev 127,
Ukraine, U.S.S.R.

The initial reason for writing this paper was to attempt a revision of the
the forms of Gonepteryx described by Prof. Dr. Shi-iti Murayama of
Ibaraki, Osaka-fu, Japan, with the aid of ultraviolet photography in
order to determine their real position within the system for the genus,
worked out in a recent monograph (Nekrutenko, 1968). In addition to
this, it will be useful to express here some taxonomic suggestions, which
I hope will show the correct position of the forms considered.

I quote Murayama (1964):

“,. . Bisher wurde aspasia aus dem Amur- und Ussuri-Gebiet als Syn-
onym von mahaguru Gistel gedeutet. Aspasia ist jedoch oberseits beim
6 am Vorderfliigel, den heller gelben Saumbereich ausgenommen, tiefer
gelb, ist grosser, und hat die Hinterfliigel weniger tief gezahnt. Daraus
ergibt sich, dass aspasia sicherlich nur eine gute Subspezies von mahaguru
sein kann.”

I offer some comments concerning this combination:

(1) Gonepteryx mahaguru aspasia of Murayama (1964) is not a new
combination. Hemming in 1935 suggested that aspasia is a subspecies of
mahaguru, taking as type locality “montagnes de Chingan jusqu’a Khokht-
sir.

(2) The question of the relationship between the names aspasia Méné-
triés and mahaguru Gistel is worthy of more detailed discussion. The
source of confusion here lies in the fact that, as was pointed out by Hem-
ming (1935), Gistel’s paper (1857 ) was unknown to his contemporaries and
was overlooked by his successors, so that the names he proposed have
never been brought into use. Search of the literature shows that there
are four principal viewpoints on this subject: (i) mahaguru and aspasia
are two names for the same species, and because the former is older than
the latter (1857 versus 1859), aspasia is to be rejected; (ii) aspasia is a
subspecies of mahaguru, which is in its turn, a senior synonym of zaneka
Moore, 1865 (Talbot, 1935); (iii) mahaguru is a subspecies of aspasia
(Hemming, 1935, et al.); (iv) mahaguru and aspasia are distinct species
(Nekrutenko, 1968). The synonymy of mahaguru to zaneka is perfectly
clear. Comparison of the texts of Gistel’s (1857) and Moore’s (1865) de-

914 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-6. Hidden wing-pattern of males of Gonepteryx spp. 1, G. mahaguru Gistel,
Kulu (Coll. Zool. Inst., Leningrad); 2, G. aspasia aspasia Mén. Amur, Radde, 22.VII.
1912. G. Kotshubey leg. (Koll. Zool. Mus. Kiev University); 3, G. a. niphonica Vty.,
Mt. Daisen, Prof. Tottori, Japan, 9.VII.1959 (Coll. Murayama); 4, G. a. iwateana
(Murayama), Paratype. Jojoji near Morioka, Pref. Iwate, Japan, 30.VII.1954 (Coll.
Murayama); 5, G. a. coreensis (Murayama), holotype, Kwangnung near Seoul, Cen-
tral Korea, 12.1X.1959 (Coll. Murayama); 6, G. a. kansuensis (Murayama), Para-
type, Kansu mer., Hsio Nganshan, 3000 m, Juli (Coll. Murayama).

scriptions show that they are both based on the same species from

Himalaya, and that both are distinct from the form described by Méne-
tries (1859).

G. aspasia and mahaguru are two separate species, and, thus all forms
described by Murayama under the specific name of mahaguru, really
belong to aspasia. Let us now consider their hidden wing-pattern (for
method and terminology see Nekrutenko, 1964, 1968 ).

Gonepteryx aspasia iwateana (Murayama, 1964)

Original description. “Kleiner als ssp. niphonica Verity. Beim 32 ist
oberseits der gelbe Farbton satter, und zieht, den Saumbereich ausgenom-

it

VOLUME 24, NUMBER 3 215

men, mehr ins Orange. Der Apex des ? springt etwas schwacher vor, der
rote Fleck am Zellschluss ist klein.”

Hidden wing-pattern (Fig. 4). Zona opaca marginalis of the forewing
narrow, far narrower than in other subspecies of aspasia. Macula lucida
centralis on the hind wing is not bright, but rather grey, and sharply
limited. This subspecies, with its orange forewing color which is well
correlated with a narrowing of the dark elements of the hidden wing
pattern, is unique within the aspasia group.

Occurrence. The subspecies was described from northern Japan
(Praef. Iwate). No additional distributional data are available.

Gonepteryx aspasia coreensis (Murayama, 1965)

Remarks. Murayama suggested (personal communication) that this
subspecies is a synonym of aspasia f. pultaina Doi (1929). Such a sugges-
tion seems questionable for the following reasons: (1) G. pultaina was
described as a spring form of aspasia, and specimens on which Murayama
based his description were collected from April to September. (2) The
main diagnostic character to distinguish pultaina are clearly visible fus-
cous spots on the underside of both wings. The latter character raises the
question as to whether pultaina is a senior synonym of f. guttifera Mell
(1943) which has very similar spots. Similar “guttiferous” aberrations were
also found among normal specimens of aspasia niphonica by Murayama
and of G. rhamni by the author. The presence of these spots may be ex-
plained, I think, by effect of environmental conditions on the overwinter-
ing insect. Thus the presence of the spots does not relate to the taxonomic
position of a given specimen. Morever, the presence of the spots has not
been pointed out in Murayama’s description (see below), nor by this
author when discussing the holotype of coreensis.

Original description. “. . . Ground colour of upperside in male more
light yellow than (in) niphonica Verity, hindwing paler yellow than in
forewing. The subspecies aspasia from Ussuri and North Korea is fairly
smaller.”

Hidden wing-pattern (Fig. 5). Zona opaca marginalis wide, wider than
in all other subspecies, occupying more than half of wing surface. Macula
lucida centralis of hindwing small, sharply limited, bright, clearly rec-
ognizable.

Occurrence. Central Korea, probably down to the southern shore of the
Korean peninsula.

Gonepteryx aspasia kansuensis (Murayama, 1965)
Original description. “The Chinese form, collected in Hsio, Nganshan,
Kansu is very allied to coreensis, but yellow ground colour of upperside in

216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

male somewhat deeper, the projecting angle of hindwing more distinctly
sharp and outer margin of that more dentate.”

Hidden wing-pattern (Fig. 6). Zona opaca marginalis of forewing some-
what wider than in niphonica, deeply penetrating into area lucida supe-
rior between veins. Macula lucida centralis of hind wing small, of nearly
same size as in coreensis.

Occurrence. Northern slope of the Kwen-lung mountain system, on the
border with Gobi desert.

Discussion and Conclusions

(1) All forms described by Murayama are valid subspecies, and the
study with ultraviolet rays supports their subspecific status within the
the species aspasia Mén.

(2) In the present state of our knowledge of variation in G. aspasia,
iwateana and coreensis appear to be extreme forms in an array of sub-
species with different degree of development of single elements of the
hidden wing-pattern.

(3) As was established for other than species of the aspasia-group of
Gonepteryx s. str., orange wing coloration is strictly correlated with reduc-
tion of the dark elements in the hidden wing-pattern. Such forms occur
under warm and wet climatic conditions. The forms considered show a
similar interrelation between visible wing color and hidden pattern in
representatives of aspasia-group, but the environmental conditions on
which these characters depend are reversed from those for the rhamni-
group. G. aspasia coreensis has far broader dark zones in its hidden wing-
pattern than aspasia aspasia from the far-eastern U.S.S.R. (Fig. 2); the
hidden wing-pattern of aspasia iwateana from northern Japan is decidedly
brighter than that of aspasia niphonica inhabiting the southern parts of
Japan. This puzzling phenomenon cannot yet be explained, because only
extensive material will provide a solution to the problem of direction of
evolutionary changes, and perhaps also provide a pattern of geographic
variation within the group.

(4) The slight hind wing dentition in aspasia kansuensis may be evi-
dence of a close relationship between aspasia and mahaguru (Fig. 1), but
once again the question arises as to which of these two species is the more
primitive.

Acknowledgments

|

| want to thank Dr. Sht-iti Murayama who kindly assisted me in this
y by lending his type specimens and providing information not con-
nis descriptions. I am grateful to Dr. Jerry A. Powell for his re-

VOLUME 24, NUMBER 3 BT

marks and criticisms, and to Dr. D. F. Hardwick who edited and cor-
rected the manuscript.

Literature Cited

Dor H., 1929. On the vernal form of Gonepteryx aspasia aspasia. J. Chosen Nat.
Hist. Soc. 8:19-20 (In Japanese, English title only).

GisteL J., 1857. Achthundert und zwanzig neue oder unbeschriebene wirbellose
Thiere (charakterisirt yon Doctor Juhannes Gistel). Straubing, Verlag der Schor-
ner schen Buchhandlung (reprint).

Hemmine F., 1935. On the identity of four species of Rhopalocera described by
Johannes Gistel in 1857. Stylops 4:121—-122.

International Code of Zoological Nomenclature adopted by the XV _ International
Congress of Zoology. International Trust for Zoological Nomenclature, London,
1964. (Russian edition: Moscow-Leningrad, 1966).

MeE.LL R., 1943. Inventur und okologisches Material zu einer Biologie der sid-
chinesischen Pieriden. Beitrage zur Fauna Sinica XXI. Zoologica (Stuttgart).
36( 100).

MENETRIES E., 1859. Lépidoptéres de la Sibérie Orientale et en particulier des
rivers de !Amour. In Dr. Leopold von Schrenck’s Reisen und Forschungen im
Amur-Lande. 2(1):17.

Moore F., 1865. List of diurnal Lepidoptera collected by Capt. A. M. Lang in the
N. W. Himalayas. Proc. Sci. Meet. Zool. Soc. Lond. p. 493.

MurayaMA S.I. 1964. Neue Tagfalterformen aus Japan und Korea. Zeitschr. Wien.
ent. Ges. 49:36.

1965. Some Korean butterflies with remarks on the related species from Japan and
China. New Entomologist 14:60-63 (In Japanese, English summary ).

NEKRUTENKO Y. P., 1964. The hidden wing-pattern of some Palearctic species of
Gonepteryx and its taxonomic value. J. Res. Lepid. 3:65-68.

1968. Phylogeny and geographical distribution of Gonepteryx ( Lepidoptera,
tera, Pieridae). Kiev, “Naukova Dumka” 130 pp., 20 pl. (in Russian, English
summary ).

Taupot G., 1935. Pieridae III. In Lepidopterorum Catalogus, ed. E. Strand, pt. 66.
W. Junk’s-Gravenhage, Berlin. pp. 518-519, 645.

NEW SPHINX MOTH RECORD FOR THE UNITED STATES

A high school student, Mr. Thane Hodson of Ottawa, Kansas found a huge moth
seated on his back porch light on August 14, 1969. Mr. William D. Field, curator of
insects at the United States National Museum, identified the moth as a male specimen
of Protoparce albiplaga (Walker). The moth was evidently a windborne example
from the neotropics. Many other tropical species such as Pholus labruscae, P. vitis,
Erynnis ello, E. alope, Erebus odora, and Thysania zenobia (see season summaries )
have all been taken at Ottawa during previous summers. However, this is the first
time that Protoparce albiplaga has been taken in the United States. It is a Brazilian
species normally occurring no farther north than southern Mexico. The specimen, in
nearly perfect condition, has been deposited in the United States National Museum.

WiiuiaAM H. Howe, 822 East Eleventh St., Ottawa, Kansas.

218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

A NEW SUBSPECIES OF GONEPTERYX RHAMNI FROM
TIAN-SHAN MOUNTAINS, U.S.S.R.

Yurt P. NEKRUTENKO
P.O. Box 324/47, Kiev 1, Ukraine, U.S.S.R.

When compiling the recent monograph of the genus Gonepteryx
(Nekrutenko, 1968 ), I felt the lack of material from areas surrounding the
great mountains of Asia, and some gaps still remain in the geographical
scope of this survey. Consideration of modest amount of material avail-
able from Tibet and the Himalayas showed this area to be a center of
subspecific diversity of Gonepteryx rhamni (L.). However, there was no
material from the neighboring mountain areas of Pamir and Tian-Shan,
so that trends in the geographic variation of G. rhamni could not be
detected.

This paper is based on material kindly collected on my request by
Dmitri S. Lastochkin of Kiev. Examination of these specimens showed
that in the Tian-Shan Mountains, or more exactly, in Alma-Ata vicinity,
occurs a population of distinctive individuals which I now describe as a
new subspecies.

Gonepteryx rhamni tianshanica Nekrutenko, new subspecies

Decidedly larger than specimens of nomenotypic race from Scandi-
navia and adjoining areas of western Siberia (alar expanse 55-60 mm
versus 40-50 mm in G. rhamni rhamni).

Male: Ground color bright, vivid, lemon-yellow; orange discal spots large, about
2 mm in diameter, easily recognizable. Hairs on thorax and abdomen whitish,
brighter than in G. rhamni rhamni. Underside of wings yellow, silky-smooth.

Superficially very close to G. rhamni transiens Verity of southern Europe.

Female: Ground color greenish, discal spots of same size and color as in male.

By hidden wing pattern (reflected ultraviolet)! this subspecies is closer to G.
rhamni rhamni than to G. rhamni transiens. Forewing zona opaca marginalis wide,
especially in anal area. Reflected elements of hidden wing-pattern on hind wing
expressed poorly, diffused, in some specimens absent.

Female showing no reflected ultraviolet, appearing dark in the photographs.

Holotype male: Union of Soviet Socialist Republics, Pravy Talgar,
Tian-Shan Mts., 1500 m, Alma-Ata prope, 8-12 July 1967 (D. S. Lastoch-
kin) (Plate, fig. 1). Allotype female: Same data (Plate, fig. 2). Para-

| Yor method of material treatment and terminology see Nekrutenko, 1964.

=>

igs. 1-6. Hidden wing-pattern of Gonepteryx rhamni tianshanica Nekrutenko;

holatyne: 9

2, allotype; 3-6, paratypes.

VoLUME 24, NUMBER 3 219

920 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

types: 1786, 1022, same data; 4646, 12 Zailijski Ala-Too, Tian-Shan
Mts., Tugok-su, 2500 m, 7 July 1968 (A. Kozubowski) (ex coll. D. S.
Lastochkin ).

All material deposited in the collection of Ukrainian Research Institute
for Plant Protection, Kiev; 26¢ and 2? paratypes forwarded to
Deutsches Entomologisches Institut, Eberswalde, D.D.R. (East Germany).

Comparative notes. As was stated, G. rhamni tianshanica occupies
an intermediate position between G. rhamni transiens and G. rhamni
rhamni. To the former it is closer by visible characters, to the second by
characters of the hidden wing pattern. Taking this into account, we
can see that G. r. transiens is a western form intermediate between G.
rhamni rhamni and G. rhamni meridionalis Rob., another described sub-
species transitional between G. rhamni rhamni, or its Siberian form, and
rhamni nepalensis Dbld. When material from the Pamir Mountains be-
comes available, the correct position of tianshanica in the system of West
Palearctic forms of G. rhamni will become even clearer.

Literature Cited

NEKRUTENKO, Y. P., 1964. The hidden wing-pattern of some Palearctic species
of Gonepteryx and its taxonomic value. J. Res. Lepid., 3: 65.
1968. Phylogeny and geographical distribution of Gonepteryx (Lepidoptera,
Pieridae), Kiev; 130 pp., 20 pl. (In Russian).

HOST-PLANT FINDING BY ODOR IN ADULT CORYPHISTA
MEADI (GEOMETRIDAE)

D. E. BreruBe!
Department of Zoology, University of Massachusetts Amherst, Mass.

Introduction

The barberry geometer, Coryphista meadi (Packard ), is a monophagous
geometrid which completes its life cycle on Berberis L. (barberry) (Dyar,
1902; Comstock, 1967). Ranging across the U. S., it is locally common
where barberry is common. The species is multivoltine and in the North-
east occurs from June through September.

by day, the moths rest among the lower barberry canes, and become
active shortly after sunset when large numbers can be observed fluttering
shove barberry shrubs. In the present study, I found that most of these

Vepartment of Biology, Yale University, New Haven, Connecticut 06520.

VOLUME 24, NUMBER 3 Din)

OLFACTOMETER

retaining screen in A
up position

One foot

Fig. 1. “Y” olfactometer. Clippings of plants to be tested are placed at A; moths to
to be tested are introduced at B; selections are determined by collecting moths at C.
Inset shows the placement of light baffles.

individuals were ovipositing females. Fluttering females were seen to
alight for a few moments, lay a single egg, and then move to a different
portion of the barberry hedge. This ovipositional behavior continued
until approximately 2200 hrs. EST, when the moths presumably moved
down among the barberry canes once again.

These behavioral observations clearly suggest that barberry foliage at-
tracts ovipositing females. Previous studies have shown that the stimuli
dictating host-plant selections in a variety of Lepidoptera are chemical in
nature (Beck, 1956; Gupta and Thorsteinson, 1960a, 1960b; Shorey, 1964).
It was suspected, therefore, that attraction of C. meadi to barberry was
the result of some volatile chemical or chemicals emanating from barberry
foliage.

Materials and Methods

Moths collected by hand-netting shortly after sunset were placed into a
Y-shaped olfactometer made of % inch plywood (Fig. 1). The olfactom-
eter was placed in a window which opened into a 10- X 13-foot room.
The window was blocked in a manner which allowed passage of air only
through the apparatus. An exhaust fan situated at the opposite end of the
room drew air slowly through the olfactometer. Recirculation of the air
was prevented by finally exhausting the air out of the room.
Holes drilled in the top of each arm of the “Y” admitted natural

999 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Choice Situation iN % Attracted to Barberry Pr

Barberry vs
Nothing

Barberry vs
Maple
TOTAL

Fig. 2. Attraction of C. meadi to the source of some chemical or combination of
chemicals emanating from barberry foliage. “Total” represents the sum of the two
choice situations. Significant deviations from a chance distribution are indicated by
asterisks: one asterisk, P < 0.05; two asterisks, P < 0.01; three asterisks, P < 0.001.

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light to the ends of the olfactometer. Two or three sprigs (three
to four inches long) of the plants to be tested were placed at either (or
both) of the ends marked “A”. Moths to be tested (7-11 per trial) were
introduced at “B”. A removable wire screen separated the moths from the
junction of the two arms of the “Y”. After the moths were introduced
into the olfactometer, the fan was started and air drawn through the
apparatus. The wire screen was kept in place for 30 minutes and was then
removed, allowing the moths access to either of the arms of the “Y”. All
trials were run for approximately five hours (0000-0500 hrs. EST). The
choices were noted by removing the covers, “C”, and collecting the moths
at both ends of the “Y”.

Two separate choice situations were tested: barberry (B. thunbergii
DC) vs. nothing, and barberry vs. red maple (Acer rubrum L.). All
tests were run for at least four consecutive nights, alternating the control
and experimental ends of the “Y” each night.

Results and Discussion

The results are summarized in Figure 2. The distributions of the moths
in the two ends of the olfactometer differed significantly from random
(by chi-square tests) in both choice situations (P < 0.05 for barberry vs.
nothing, and P < 0.01 for barberry vs. maple. Statistical analysis of the

left-right distribution of the moths showed no significant deviations from
random (P > 0,30). The results clearly demonstrate that C. meadi may be
attracted by a volatile substance (or substances) emanating from bar-

herry and ;
DCTTY iid |

capable of orienting up-wind and flying along a concentra-

VOLUME 24, NUMBER 3 De

tion gradient to the source of this attractant. These experimental data
support the suggestion from field observations that ovipositing females are
attracted to the host plant by olfaction. More interesting, however, these
data indicate that C. meadi may have the ability to locate its host plant
over comparatively long distances. The ovipositional behavior of the
species (flight between the laying of each egg) perhaps serves to reduce
intraspecific competition by effecting greater larval dispersal. If this is
true, then the ability to locate the host plant over relatively long distances
would apparently be highly adaptive because barberry under natural
conditions does not normally grow in dense thickets but is scattered in
patches.

This study has also indicated that the problems encountered in at-
tempting to investigate host-plant selections using olfactometers (Thor-
steinson, 1960) can be overcome. Early designs of the apparatus used in
this study did not contain the light baffles shown in Fig. 1. The moths
failed to make selections in these devices, apparently showing only posi-
tive phototropic responses. When all light was excluded from the ap-
paratus, the moths merely sat in the box where they were introduced. By
adjusting the light intensity entering the ends of the arms, responses to the
plant clippings could be obtained. The introduction of a retaining screen
further improved the responses obtained. Without the screen many of
the moths introduced into the olfactometer immediately flew, at random,
to either of the two ends of the “Y’. This was apparently an escape re-
sponse which was thwarted by keeping the screen in place for approxi-
mately 30 minutes.

Acknowledgments

I wish to thank the Massachusetts Audubon Society for granting per-
mission to use Arcadia Wildlife Sanctuary, Easthampton, Massachusetts
for collecting the moths used in this study. I would also like to thank Dr.
T. D. Sargent who provided helpful criticism in all phases of this work.

Literature Cited

Beck, S. D. 1956. The European corn borer, Pyraustra nubilalis (Hubn.), and its
principal host plant, I. Orientation and feeding behavior of the larva on the
corn plant. Ann. Ent. Soc. Amer. 49: 552-558.

Comstock, J. A. 1967. Notes on the early stages of the barberry geometrid moths,
genus Coryphista, and the description of a new subspecies of C. meadii (Lepidop-
tera). Bull. So. Calif. Acad. Sci. 66: 92-98.

Dyar, H. G. 1902. Life histories of North American Geometridae, XXXV. Psyche
9: 396.

Gupta, P. D. AND A. J. THorsTEINSON. 1960a. Food plant relations of the diamond-
backed moth (Plutella maculipennis Curt.) I. Gustation and olfaction in relation
to botanical specificity of the larvae. Ent. Expl. Appl. 3: 241-250.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bo
bo
aN

. 1960b. Food plant relations of the diamond-backed moth ( Plutella
maculipennis Curt.) II. Sensory regulation of oviposition of the adult female.
Ent. Expl. Appl. 3: 305-314.

SHorey, H. H. 1964. Sex pheromones of noctuid moths. II. Mating behavior of
Trichoplusia ni (Lepidoptera: Noctuidae) with special reference to the role of
the sex pheromone. Ann. Ent. Soc. Amer. 57: 371-377.

THorsTemnson, A. J. 1960. Host plant selection in phytophagous insects. Ann. Rev.
Ent. 5: 193-218.

A BILATERAL GYNANDROMORPH OF PIERIS RAPAE
(PIERIDAE)

A bilateral gynandromorph of the Imported Cabbage Butterfly, Pieris rapae Lin-
naeus, was taken September 18, 1969 at the Ithaca, New York city dump (Tompkins
Co.). It turned up in a large, randomly collected sample of P. rapae taken for other
purposes, and was not recognized as a gynandromorph until already dead. There is
thus no information available on its behavior. Nearly all of the butterflies in the
sample were taken in flight or on blossoms of Great Burdock (Arctium lappa Lin-
naeus ).

The specimen is female on the left side and male on the right, similar to one taken
in Bedfordshire, England in 1938 (S. H. Kershaw, Proc. S. Lond. ent. nat. Hist. Soc.
1954-55, p. 33). All secondary sexual color and pattern characteristics, including
the pteridine pigments, are normally developed on the half of appropriate sex. There
is no irregular mosaicism. The external genitalia are also bilaterally asymmetrical,
with a somewhat aborted clasper on the male side. The internal anatomy was not
studied.

A quick survey of the British aberrational literature suggests that gynandromorphs
are much rarer in P. rapae than in at least some populations of the P. napi complex.
Bilateral gynandromorphs also seem to be very rare in P. brassicae Linnaeus, though
irregular mosaics are rather frequent. My specimen is the first of its kind to appear
among perhaps 75,000 wild and bred Pieris I have examined in recent years, in-
cluding some 12,000 wild P. rapae from the northeastern United States.

Artuur M. SHAPIRO, Department of Entomology and Limnology, Cornell University,
Ithaca, New York.

A NOTE ON THE SYSTEMATIC POSITION OF PAPILIO
ANTIMACHUS

PAuL R. EHRLICH

Department of Biological Sciences, Stanford University, Stanford, California

lkecently, the Honorable Miriam Rothschild and Professors J. von Euw,
and ‘T. Reichstein (1970) have been able to prove the presence of car-
denolide heart poisons in Papilio antimachus Drury. P. antimachus is very
unusual in its appearance, with very long wings and a pattern which makes
it look something like a giant Acraea. The unusual appearance of the
putterfly, and the presence of the cardenolides (heart poisons which also

VOLUME 24, NUMBER 3 225

are called cardiac glycosides ), led Dr. Rothschild to write to me concern-
ing the systematic position of antimachus. I had not been able to examine
a specimen of this unusual butterfly in the course of previous work on the
higher classification of the Papilionoidea (Ehrlich, 1958; Ehrlich and
Ehrlich, 1967), but Dr. Rothschild was able to arrange the loan of a
specimen for dissection from the British Museum, through the courtesy of
Dr. T. G. Howarth.

Papilio antimachus turns out to be a quite typical member of the family
Papilionidae, subfamily Papilioninae. It conforms entirely to the diagnosis
of the family (Ehrlich, 1958, pp. 336-337 ) with the exception of characters
number 42 and 43 for which no information is available. Similarly, it
agrees entirely with the subfamily diagnosis (pp. 338-339) except for
character P on which no information is available. Further information on
the family characters is as follows: 6, antennae close together; 11, anterior
tentorial arms bearing very small crest; 13, cervical sclerites united beneath
neck by unbroken sclerotic band; 15, spinasternum produced in much the
manner of that of Papilio machaon (Fig. 34 of Ehrlich, 1958) except that
areas of sclerotization heavier; 19, patagia membranous; 27, mesothoracic
anepisternum not a separate sclerite; 32, caudal part of metathoracic epi-
meron broad; 34, postspiracular bar absent. Additional information on
subfamily characters is as follows: H, prodiscrimen represented by a
prominent anterior spine; L, lamella of metadiscrimen curved downward
to base of furca; O, tarsal claws symmetrical and simple.

In the most recent treatment of the tribes of the Papilionidae (Ehrlich
and Munroe, 1960) P. antimachus falls into the tribe Papilionini. At the
moment the placement of antimachus by Munroe (1961) in the genus
Papilio seems entirely appropriate, although examination of the early
stages may require revision of the generic and tribal position.

Literature Cited

Eueticy, P. R., 1958. The comparative morphology, phylogeny and higher classifica-
tion of the butterflies (Lepidoptera: Papilionoidea). University of Kansas
Science Bulletin 39: 305-370.

Eureuicu, P. R., anp A. H. Enruicu, 1967. The phenetic relationships of the butter-
flies. J. Adult taxonomy and the nonspecificity hypothesis. Systematic Zoology
16: 301-317.

Munroe, E., 1961. The classification of the Papilionidae (Lepidoptera). Canadian
Entomologist, Suppl. 17, 51 pp.

Muwnrog, E., AND P. R. Enruicnu, 1960. Harmonization of concepts of higher classifi-
cation of the Papilionidae. Journal of Lepidopterists’ Society 14(3): 169-175.

RotTHscHILD, M., J. von Euw AnpD T. REICHSTEIN, 1970. Cardenolides in Papilio anti-
machus Drury (Papilionidae) together with records of Lepidoptera feeding on
Asclepiadaceae or Apocynaceae which do not store heart poisons in their body
tissue. Ms. in preparation.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bo
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FURTHER EASTWARD EXTENSION OF THE RANGE OF
EUCHLOE AUSONIDES MAYI (PIERIDAE) IN ONTARIO

Euchloe ausonides mayi Chermock & Chermock, is first known to have been
collected in Ontario near the Manitoba border in 1947. Riotte (1968, J. Lepid.
Soc., 22: 40) has traced the eastward extension of its range which reached
Geraldton by at least 1966. In this locality he had collected extensively until 1958
without seeing it. In 1965 I collected the species on Manitoulin Island, Ontario, 330
miles southwest of Geraldton for the first time in spring. That year, and again in
1966, 1967 and 1968, this species first appeared during warm weather in the second
week of May, remaining common until the month’s end. No June collecting has
been done, and the species has never been seen there during July. All records are
from the limestone barrens in Burpee Township near the shore of Lake Huron,
where the insect flies close to ground in sunny clearings in the juniper-white cedar-
white spruce—poplar scrub known locally as “prairies.” Arabis drummondi, a known
host, is frequent in this habitat. Identification of the butterflies has been confirmed
by J. C. E. Riotte, and two specimens taken May 13, 1968 have been placed in the
collection of the Royal Ontario Museum, Toronto. It may be that the species, which
is now common around Fort William, Ontario, proceeded along the north shore of
Lake Superior to the east and thence to Manitoulin Island.

RONALD R. TAsker, 12 Cluny Dr., Toronto 5, Ontario.

COURTSHIP AND MATING BETWEEN CHLOSYNE NEUMOEGENI
AND CHLOSYNE CALIFORNICA (NYMPHALIDAE)

The ranges of Chlosyne neumogeni (Skinner) and Chlosyne californica (Wright)
overlap in certain areas of southern California. In many of the desolate mountain
ranges of the Mojave Desert both species can be found flying together. While on a
collecting trip to the Providence Mountains, I discovered that both C. neumogeni and
C. californica were breeding in the area. After hiking out of Bonanza King Mine
Canyon on April 20, 1969, I noticed large numbers of these two species of butterflies
fluttering around flowering bushes. It was at the entrance of this canyon, at an eleva-
tion of approximately 4,000 ft., that I observed courtship between these two species.
I saw battered C. neumoegeni males chasing C. californica females in the air. When
these females alighted upon flowers, with spread wings, the males would hover a
couple of inches above them. Later that afternoon, I began hiking towards Gilroy
Canyon, about a mile south of Bonanza King Mine Canyon. Climbing down the side
of a ravine into a dry streambed, I spotted two mating Chlosyne resting together on a
flowering bush. Upon closer observation, I noticed that the two butterflies in copula-
tion were a male of C. newmoegeni and a female of C. californica.

No examples of courtship and mating between male C. californica and female C.
neumoegeni were observed in the Providence Mountains at this time. Also, C. califor-
nica appeared to be approximately eight times more common than C. neumoegeni in
this region. C. californica seemed to be at the peak of its flight period, because more
than half of the specimens captured were fresh. C. neumoegeni seemed to be in the
latter part of its flight period, because most specimens were worn. One wonders if
these factors might be partially causative of this rare phenomena of courtship and
mating between two distinct species.

RicHARD C, Priestar, 5631 Cielo Ave., Goleta, Calif.

VOLUME 24, NUMBER 3 eT

A NEW FOODPLANT RECORD FOR ECPANTHERIA SCRIBONIA
(ARCTIIDAE )

During a collecting trip to Florida in 1968 I found a larva of Ecpantheria scribonia
(Stoll) feeding on the underside of a fern, Polypodium aureum L., 2 miles southwest
of Satsuma, Putnam County, Florida. This larva, when collected cn 31 June 1968,
was in the penultimate instar. It pupated on July 26th and emerged, as a female,
on 14 August 1968. The host fern is epiphytic on trees, especially palmetto, and
has a subtropical and tropical distribution. A search of the literature has proven
this a new hostplant record, and apparently is also the first time any fern has been
recorded as a foodplant for this moth. Sidney A. Hessel kindly assisted with the
identification of the fern.

As a postscript, I would like to add that I was brought a last instar larva of E.
scribonia collected by Eugene S. Morton at Leete’s Island, Guilford, New Haven
County, Connecticut, on 29 October 1968. After some prolonged feeding on
dandelion and plantain, it pupated; an adult female emerged on 13 December 1968.
There seem to be no previous records of larvae in Connecticut. E. scribonia occurs
further north, at least to Massachusetts and southern Ontario, but is rare in
Connecticut.

Both specimens have been deposited in the Peabody Museum of Natural History,
Yale University.

JoszepH A. ConcELLo Jr., Peabody Museum of Natural History, Yale University,
New Haven, Connecticut.

CRAMBIDIA PURA (ARCTIIDAE: LITHOSIINAE) NEW FOR CANADA

During the summer 1969 two lepidopterological teams were working at the Biologi-
cal Station of Queen’s University, Kingston, Ontario, in Chaffeys Locks, Leeds Co.,
Ontario. One team was from the Royal Ontario Museum, the other from the Dept.
of Biology of Queen’s University (Prof. R. Harmsen). Both teams worked together in
identifying and coordinating the catches from six “Black Lights,” of which five were
located at the station and one at Glenburnie, Frontenac Co., Ontario. The lights at
the station were in rich deciduous forest, the light in Glenburnie in normal farming
country.

In past years the lights at the station had attracted a few rare Lithosiinae, like
Cisthene unifascia Gr. & Rob. This year we took for the first time at the station
Crambidia casta Packard which was known to occur in this part of the country.

In Glenburnie, however, numerous specimens of Crambidia pura Barnes & Mc-
Dunnough, a species never before recorded from Canada were taken. It was described
from North Carolina, and Forbes (1960, Lepidoptera of New York and neighboring
states, pt. IV. Cornell Univ. Agr. Exp. Sta. Mem. 371) gave its range as Minetto,
New York; Lakehurst, New Jersey, and North Carolina.

In Glenburnie Crambidia pura, the identity of which was ascertained by genitalic
dissection, would seem to have two generations, one at the end of June, the other in
the middle of August.

J. C. E. Riorre, Research Associate, Department of Entomology, Royal Ontario
Museum, Toronto, Ontario, Canada.

928 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NOTES ON CALLOPHRYS (CALLOPHRYS) COMSTOCKI HENNE
IN NEVADA

According to Ehrlich and Ehrlich (1961, How to know the Butterflies), Callophrys
(Callophrys) comstocki Henne was known only from the Providence Mountains of San
Bernardino Co., California. Thus, the discovery of this species in Nevada is an impor-
tant extension of its known range. This finding was made on March 23, 1968 when
Kenneth Tidwell and I were on a collecting trip in Nevada. At the time, we were
collecting in the dry hills about five miles west of Goodsprings, Clark County, Nevada,
just north of Wilson Pass, at about 5000’. At about 11:00 A.M., when Ken and I were
exploring a large arroyo near a radio relay station, I observed a small butterfly flying
in a rocky side canyon on the east facing slope of the arroyo. Though at first thought
to be a skipper, closer observation revealed it to be a species of Callophrys. It eventually
rested on a rock in the dry stream bed and was captured. Further specimens were
observed sitting on rocks or on the sand. They darted up when disturbed, but would
shortly return to the same spot after circling around a few times, in skipper fashion,
before alighting. They showed no preference for any particular kind of vegetation. A
total of 3 females and 5 males, all in fresh condition, were captured.

One specimen was later examined by Harry K. Clench who determined it to be
Callophrys (Callophrys) comstocki Henne. As compared with Providence Mountains
material, it was found to be slightly larger with darker black markings on the post-
median line of the underside.

In the same small canyon as the comstocki were taken, were found Callophrys fotis
fotis (Strecker) and Callophrys siva (Edwards). The C. fotis were about as common
as the comstocki, but preferred resting on bushes instead of the ground. A single C.
siva was captured while sitting on a stunted juniper bush, a few of which grew along
the sides of the canyon. The three aforementioned species appeared to be quite local
as Ken Tidwell was unable to locate additional specimens in the surrounding area.

On the ridge above the canyon, many butterflies were found “hilltopping.” Among
these were Papilio rudkini, Anthocaris sara inghami Gunder, Euchloe creusa (Double-
day), and Pieris sisymbri Boisduval.

Curtis J. CALLAGHAN, 2500 Kensington Ave., Salt Lake City, Ut.

BOOK REVIEW

A sHort History OF THE BROWN-TaiL Motu, by William Curtis. A Curwen facsimile
edition, 1969; 18 + 12 pp., 1 coloured plate. Introductions by W. T. Stearn and
D. S. Fletcher. Distributed in North America by Entomological Reprint Specialists,
P.O. Box 207, East Lansing, Michigan. Price $9.30 U. S.

This is a beautifully prepared little book that will be of particular interest to the
bibliophile. The edition has been limited to a run of 1,000 copies. In the facsimile
reproduction of William Curtis’s classical treatment of the Brown-Tail Moth, the
history of an outbreak, the seasonal development of the insect, its food plants and
its potential predators are discussed from the stand-point of a biologist of 1782. A
biographical introduction by W. T. Stearn outlines what is known of the life of
William Curtis (1746-1799), primarily a botanist but latterly an entomologist. An-
other introductory section by D. S. Fletcher of the British Museum (Natural History)
is devoted to the taxonomic relationships of the Brown-Tail, its pest status as a forest

sect, and the history of its introduction and spread in North America.
» Harpwick, Editor.

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NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
for material in excess of 20 printed pages, at the rate of $17.50 per page. Address
all correspondence relating to the Journal to: Dr. D. F. Hardwick, K. W. Neatby
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Contributors should prepare manuscripts according to the following instructions;
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Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
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full scientific name, with authors of zoological names. Underline only where italics
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numbered consecutively, and the footnotes should be typed on a separate sheet.

Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
heading LirerRatTurE CitTep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: All photographs and drawings should be mounted on stiff, white
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Material not intended for permanent record, such as current events and _ notices,
should be sent to the editor of the News: Dr. C. V. Covell, Dept. of Biology, Univer-
sity of Louisville, Louisville, Kentucky 40208.

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyrit F. pos Passos
Price, postpaid: Society members—$4.50, others—$6.00; uncut,
unbound signatures available for interleaving and private binding,

same prices; hard cover bound, add $1.50. Revised lists of the
Melitaeinae and Lycaenidae will be distributed to purchasers free.

ALLEN PRESS, INC. ake LAWRENCE, KANSAS
US. me

CONTENTS

Berube, D. E. Host-plant finding by odor in adult Coryphista meadi ( Geo-

metridae) (2:0
Callaghan, C. J. Notes on Callophrys (Callophrys) comstocki Henne in
Nevada 22.2 A
Concello, J. A. A new foodplant record for Ecpantheria scribonia ( Are-
tlidae )

Doyle, J. F. III. Field notes on three skippers in Texas (Hesperiidae) _____
Ehrlich, P. R. A note on the systematic position of Papilio antimachus ___
Ferris, C. D. Occurrence of Coenonympha inornata (Satyridae) in Maine

Hardwick, D. F. Book Review: A short history of the Brown-Tail Moth,
by William ‘Curtis: 20s 0 ONE Oe

Howe, W. H. New sphinx moth record for the United States _____________

Leech, H. B. The aegeriid Ramosia fragariae in a flight trap, and the in-
terpretation thereof i092 Ue

Malicky, H. New aspects on the association between lycaenid larvae
(Lycaenidae) and ants (Formicidae, Hymenoptera)

Masters, J. H. Distributional notes on the genus Mestra (Nymphalidae)
in North Ameériea (0) Ge ON Sa

Mather, B. Variation of Graphium marcellus in Mississippi (Papilionidae )

Nekrutenko, Y. P. Comments on forms of Gonepteryx aspasia (Pieridae )
described by Shu-iti Murayama eee

Nekrutenko, Y. P. A new subspecies of Gonepteryx rhamni from Tian-Shan
Mountains, )\U:.S.SiR (i a i) rr

Priestaf, R. C. Courtship and mating between Chlosyne neumoegeni and
Chlosyne californica (Nymphalidae) _.....

Rindge, F. H. Freeman collection of Megathymidae donated to the
American Museum of Natural History

Riotte, J. C. E. Crambidia pura (Arctiidae) new for Canada

Shapiro, A. M. A bilateral gynadromorph of Pieris rapae (Pieridae)

Tasker, R. R. Further eastward extension of the range of Euchloe ausonides
mayt (Pieridae) in Ontario 0 ee
Williams, C. B. The migrations of the painted lady butterfly, Vanessa cardui
(Nymphalidae ), with special reference to North America

Wright, D. A. The effect of photoperiod on the termination of pupal
diapause in the wild silkworm, Actias luna

212
224-225
202

228
217

189

190-202

203-208
176-189

213-217

218-220

226

208
22:7
224

226

157-175

209-212

\

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Volume

9A. 1970 Number 4

JOURNAL

of the

_ Lepivoprerists SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Be chen von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

16 November 1970

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

D. F. Harpwick, Editor of the Journal
C. V. CovE.u, Editor of the News
S. A. HesseL, Manager of the Memoirs

EXECUTIVE COUNCIL

E. B. Forp (Oxford, England), President

C. L. Remineron (New Haven, Conn. ), President-elect
L. P. Brower (Amherst, Mass.), Ist Vice President

L. M. Martin (Prescott, Ariz.), Vice President

J. W. TrwvEN (San Jose, Calif.), Vice President

S. S. Nicouay (Virginia Beach, Va.), Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

Members at large (three year term): W. C. McGurrin (Ottawa, Ont.) 1971
J. F. G. Crarxe (Washington, D.C.) 1970 Y. Nexrurenxo (Kiev, U.S.S.R.) 1971
H. K. Cuencu (Pittsburgh, Penna.) 1970 B. Marner (Clinton, Miss.) 1972

B. Wricut (Halifax, N.S.) 1970 M. Ocata (Osaka, Japan) 1972

A. E. Brower (Augusta, Me.) 1971 E. C. Wextuine (Merida, Mexico) 1972

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
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JOURNAL OF

Tue LepiporrerRiIstTs’ SOCIETY

Volume 24 1970 Number 4

IS PAPILIO GOTHICA (PAPILIONIDAE) A GOOD SPECIES

C. A. CLARKE AND P. M. SHEPPARD

Department of Medicine and Department of Genetics,
University of Liverpool, England

Remington (1968) has named the Papilio zelicaon-like swallowtail
butterflies from a restricted geographical range (the high mountains of
New Mexico, Colorado, and Wyoming) Papilio gothica Remington. Since
the criteria used by Remington for claiming the existence of this newly
named species are chiefly genetical and ecological rather than the usually
used morphological and behavioural ones, it seems desirable to examine
the genetic evidence more critically than Remington appears to have
done.

Genetical Evidence Obtained by Hybridization Experiments

Remington showed that P. zelicaon Lucas and P. gothica are morpho-
logically very similar and he also indicated that a number of specimens
cannot be classified unless the place of their origin is known. However,
the F, hybrids between P. gothica and P. polyxenes Fabr. on the one
hand, and P. zelicaon and P. polyxenes on the other, are distinguishable,
as are the F; hybrids when P. bairdi is substituted for P. polyxenes.

P. polyxenes and P. bairdi Edwards are much blacker insects than P.
zelicaon. They show a great reduction in the amount of yellow on both
wings and body. Clarke and Sheppard (1955, 1956) have demonstrated
that the marked difference between the color patterns of P. polyxenes
and P. zelicaon and, in fact, between P. polyxenes and the yellow Euro-
pean species, Papilio machaon L., is due to the presence of a single
major gene which is dominant or nearly dominant in effect. The P.
gothica x P. polyxenes hybrids differ from the P. zelicaon x P. polyxenes
hybrids in that those involving P. gothica have a reduction in the yellow
markings on the upper side of the wings and on the abdomen, even
more marked than in the P. zelicaon hybrids (Remington, 1968). Con-
sequently, it can be concluded that the P. gothica insects that have been

230 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Male offspring from the mating P. polyxenes 2 x P. machaon 6. Three
offspring are from a single mating, the fourth (top left) is from a second mating.
Note that even full brothers can differ more from one another than the polyxenes x
gothica and polyxenes < zelicaon illustrated by Remington.

tested contain different modifiers of the effect of the major dominant
allelomorph of P. polyxenes from those of P. zelicaon. However, genetic
work that has been undertaken in the Lepidoptera (Ford, 1955b; Clarke
and Sheppard, 1963, 1968; Sheppard, 1969) shows that different popula-
tions of a single species and even different individuals within a popula-
tion often contain modifiers having a much more marked effect than
those reported by Remington. Furthermore, the F, hybrids illustrated
by Remington show differences no more extreme and of exactly the same
type as those found by Clarke and Sheppard (1955) between individuals
when they hybridized P. polyxenes and P. machaon (Fig. 1). Thus,
‘here is good reason to believe that the differences between the F; hybrids
reported by Remington are no greater than those usually found within
a single species. Furthermore, Remington’s data do not suggest that the
‘ferences necessarily apply to all populations which he has designated
composed of P. gothica but only to those from a limited area.

VoLUME 24, NuMBER 4 OM

Sex Ratio in Hybrids

Remington also supported his argument by claiming that P. gothica
and P. zelicaon “have different hybrid sex ratios in their crosses with
P. polyxenes, gothica < polyxenes being nearly lethal for the sex of the
polyxenes parent whereas zelicaon X polyxenes had more nearly equal
sex ratio although deficient for the sex of the polyxenes parent.” Even
if Remington’s claim were validated (it is not supported by his data,
the statistical procedures he used being wrong) the mere fact that sex
ratios can be profoundly different in reciprocal hybrid matings (Haldane,
1922) should warn the unwary that very small genetic differences may
profoundly alter sex ratios in hybrids. Furthermore, even a cursory
knowledge of Lepidopteran genetics would acquaint one with the fact
that very different sex ratios may appear even between the progeny of
sibs of the same sex when hybridization is involved. In fact, the extreme
sensitivity of sex ratio to minor genetic differences in the parents is
demonstrated by Remington’s own data. Thus, the two crosses he re-
ports between female zelicaon < male polyxenes show significantly dif-
ferent sex ratios among the progeny (P = 0.004, Fisher's exact test). It
is because of this heterogeneity that it is illegitimate to combine the two
broods as Remington has done in his Table 1B for comparison with the
single cross of a female gothica X male polyxenes. If the correct statistical
procedure is followed we find that one of the two zelicaon hybrid broods
is significantly different from the gothica brood (P = 0.0007) but the
other is not (P = 0.122). Thus, there is no evidence as yet that the two
types of hybrid (those involving zelicaon and those involving gothica)
generate different sex ratios. Even if such evidence is eventually forth-
coming this would not indicate that the two forms are genetically very
dissimilar.

Polymorphism for Larval Spot Color

Remington noted from his limited experience of wild P. zelicaon larvae
(less than 50 independent observations) that they are polymorphic for
the color of their sub-dorsal spotting. Previously, we had shown (Clarke
and Sheppard, 1955, 1956) that this polymorphism is mainly controlled
by a single gene. More extensive sampling in Napa, Yolo and Eldorado
Counties (Sheppard, unpublished) is in agreement with Remington's
view that the polymorphism is widespread in lowland Californian popu-
lations of P. zelicaon. On the other hand, in all of about 20 independent
observations (Remington, 1968) the larvae of P. gothica were mono-
morphic, being homozygous for the allelomorph producing yellow spot-
ting. Remington suggests that in this respect they are unusual. However,

232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

although P. polyxenes is polymorphic for red spotting in some popula-
tions, as Remington states, the allelomorph responsible is often absent
or at such a low frequency in P. polyxenes populations that these cannot
be considered to be different from those of P. gothica with respect to
the polymorphism on the available evidence. The gene frequencies in
the populations of P. zelicaon and P. gothica so far reported are clearly
different. However, since P. gothica has merely become monomorphic
for a character polymorphic in other areas the difference does not in-
dicate any profound genetic change. Parallel examples are common in
the Lepidoptera in situations where there is no question of two different
species being involved (Clarke and Sheppard, 1963, 1968). Even within
the machaon group itself in North America the different frequencies of
the forms comstocki and clarki in populations of P. rudkini Comstock
may illustrate the point.

Ecological Evidence

P. gothica is strictly univoltine in Colorado, whereas P. zelicaon, as
well as many other members of the machaon group, is multivoltine. This
genetic difference has not yet been analyzed in detail. However, the
difference is not surprising since P. gothica is from the high mountains
where a single generation a year would seem to be ecologically advan-
tageous. In fact, the difference from P. zelicaon merely suggests that
both are ecologically adapted to their respective environments. Thus
there are two ecotypes, as is so often found when montane or northern
and lowland or southern types of a single species are compared. Ex-
amples from Lepidoptera in the British Isles which illustrate this point
are the butterflies Aricia agestis Schiff., and Polyommatus icarus Rott.
(Ford, 1945) and the moth Lasiocampa quercus L. (Ford, 1955a), the
latter having races with a two year and a one year life cycle.

P. gothica also differs from P. zelicaon in that the larvae of the former
appear to be found in nature only on Pseudocymopterus montanus (A.
Gray ) Coulter and Rose (five larvae and an unknown number of eggs
reported by Remington, 1968) whereas several species of Umbelliferae
are utilized by P. zelicaon. However, this restriction of P. gothica but
not P. zelicaon to a single larval food plant, even if it is substantiated
by more evidence, would not indicate that P. zelicaon and P. gothica are
different species, for such ecotypic variation is common in the Lepi-
doptera and is found even within the machaon group itself. Thus, the
P. machaon race from eastern England not only shows marked behay-
ioural differences compared with the continental race but the larvae are
confined to the milk parsley, Peucedanum palustri L., whereas the con-

VoLUME 24, NuMBER 4 233

tinental form is not so confined (Ford, 1945). This is a striking parallel
within a single species to the difference reported by Remington.

All the genetic evidence presented by Remington suggests that gothica
is only a minor high mountain ecotype of P. zelicaon and is not a good
species in its own right. Unless evidence can be produced to show that
the two forms are sexually isolated if they come together in nature (a
matter that should not be difficult to study experimentally), then the
use of gothica as a specific name should be discontinued. Its continued
use will only confuse and not clarify the taxonomy of the machaon group.
Although genetic evidence is valuable in taxonomy it must not be in-
terpreted in a parochial way. Only if it is considered in the light of
studies on other material from other lands will it be useful in clarifying
taxonomic relationships. We fully agree with Remington’s (1968) state-
ment that the machaon group “is much too complicated for grand con-
clusions based on scanty breeding experiments or on specimen samples
from a few distant, randomly-chosen localities.”

Literature Cited

CLARKE, C. A. AND P. M. SHepparp. 1955. A preliminary report on the genetics
of the machaon group of swallowtail butterflies. Evolution 9: 182-201.
1956. A further report on the genetics of the machaon group of swallow-

tail butterflies. Evolution 10: 66-73.

1963. Interactions between major genes and polygenes in the determina-

tion of mimetic patterns of Papilio dardanus. Evolution 17: 404-413.

1968. The genetics of the mimetic butterfly Papilio memnon L. Phil.
Trans. Roy. Soc. 254: 37-89.

Forp, E. B. 1945. Butterflies. Collins, London.

1955a. Moths. Collins, London.

1955b. Polymorphism and taxonomy. Heredity 9: 255-264.

Hapang, J. B. S. 1922. Sex ratio and unisexual sterility in hybrid animals. J.
Genet. 12: 101-109.

ReminctTon, C. L. 1968. A new sibling Papilio from the Rocky Mountains, with
genetic and biological notes (Insecta, Lepidoptera). Postilla 119: 1-40.

SHEPPARD, P. M. 1969. Evolutionary genetics of animal populations: the study of
natural populations. Proc. 12th Int. Cong. Genet. 3: 261-279.

A NEW RECORD FOR NORTH AMERICA OF A SWALLOWTAIL BUTTERFLY
(PAPILIONIDAE )

While visiting with Mr. J. E. Lipes in El Salvador, I was privileged to examine
some of the material collected by him. Among his material were three male specimens
of Papilio (Graphium) philolaus Bsd. Mr. Lipes informed me that all three were taken
in Texas. However, only one specimen had complete data, which are as follows: July
21, 1958, Padre Island, Port Isabel, Cameron County, Texas, leg. J. E. Lipes. The
specimen is being donated to the Allyn Foundation collection, Sarasota, Florida.

RAYMOND J. JAE, 1286 South Umatilla Street, Denver, Colorado.

934 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

THE LIFE HISTORY OF PYRRHIA EXPRIMENS (NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario

In more recent North American literature, the genus Pyrrhia has been
included in the noctuid subfamily Amphipyrinae; genitalic characters
show it to be a true constituent of the subfamily Heliothidinae, however.
This relationship is substantiated by the larval habit of feeding primarily
on the flowers and fruits of its host plants, and by the structure of the
larva itself which is close to that of other species of the Heliothidinae
(Crumb, 1956; Gardner, 1946).

There are two North American species of Pyrrhia: umbra (Hufn.) and
exprimens (Wlk.). The latter has, until recently, been considered only a
subspecies of umbra. For much of their respective distributions, how-
ever, the two occur sympatrically, and they differ not only in maculation
and colouring but in the structure of the male genitalia. In exprimens the
ground colour is orange-brown rather than golden as in umbra; in the male
genitalia, both the valve and the ampulla are shorter in exprimens than in
umbra.

In eastern North America, exprimens is generally of more northern dis-
tribution than is umbra. The former is distributed in southern Canada
from the Atlantic to Pacific coasts. In eastern North America, it occurs
southward at least to Pennsylvania, and in the west at least to Colorado.
The few specimens I have from the latter state are all of the heavily suf-
fused form, stilla Grt., and these may represent a distinctive southwestern
race of exprimens.

Pyrrhia exprimens is rather a general feeder, being recorded from a wide
variety of plants. Forbes (1954) listed larvae as feeding on Polygonum,
Desmodium, and Juglans. The Canadian Forest Insect Survey (Prentice,
1962) has recovered larvae from Populus, Salix, Rosa, and Prunus. In
addition to these records, there are specimens in the Canadian National
Collection reared from Althea, Betula, and Delphinium. The species is,
on occasion, a pest of delphinium in the Ottawa area, the young larvae
teeding on the blossoms and boring through the unopened buds.

A BRA ter ° 3 0 Ono

Although exprimens is obviously broadly polyphagous, the ovipositing
‘female does seem to show definite preferences in host plant selection. A
single female of exprimens was exposed to four sprays of plants in an ovi-

position container. Each of the sprays was of comparable size, and each
vas replaced daily in the oviposition container. Over a period of eleven
cl i | an oh | | eae b ’ . “4 °

ne numbers of eggs laid on each were as follows: Geranium sp.,

VOLUME 24, NuMBER 4 Zoo

447; Phlox sp., 151; Chrysanthemum sp., 64; Ranunculus sp., 33. In addi-
tion to these, 29 eggs were laid on the wads of cotton wool that secured
the sprays in vials of water. Eggs were deposited singly at the apex of the
food plant. In all, the single wild-caught female deposited 724 eggs.

Pyrrhia exprimens overwinters as a pupa in a cell a few inches below the
surface of the ground. On the basis of records available, the species is
single-brooded throughout its range.

Descriptions of Stages

The descriptions of immature stages in the following section were based
on the progeny of a single female taken at Ottawa, Ontario. The rearing
methods employed were those outlined by Hardwick (1958). Larvae
were reared individually on cranesbill (Geranium sp.) at room tempera-
ture. Of 63 larvae reared to the pupal stage, 21 matured in five larval
stadia, 42 matured in six stadia, and one matured in seven stadia. The
estimate of variability, following the means for various measurements, is
the standard deviation.

Adult (Figs. 1, 2). Ground colour orange-brown rather than golden as in wmbra.
Head and thorax clothed with mixed spatulated and hair-like vestiture. Head and
thorax orange-brown, concolorous with forewing; abdomen predominantly greyish-
brown dorsally. Forewing orange-brown, variably suffused with violet or reddish-
brown in subterminal space and outer half of median space. Transverse anterior line
forming three distinct outward angles. The anterior half of a basal line usually
evident. Transverse posterior line weakly bisinuate, angling strongly inward from
costal to trailing margins. Orbicular spot large, usually circular, occasionally with
some dark central shading. Reniform spot large, irregular, usually darkly suffused
centrally. Median line well defined, angling outward to postero-inner angle of reni-
form, then angling inward to trailing margin. Median space, distal to median line,
variably suffused with violet or reddish-brown, occasionally heavily so. Subterminal
line irregular, scalloped between the veins. Subterminal space usually suffused with
violet or reddish-brown. Terminal space concolorous with basal area of median space.
Usually a narrow, dark, terminal line. Fringe concolorous with terminal space. Hind
wing pallid yellow, with a rather broad, chocolate-brown, outer-marginal band. A
prominent brown discal lunule. A rather diffuse brown band along inner margin.
Fringe pale yellow. Underside of forewing rather dark yellow with a slender, elongate,
chocolate-brown basal dash, a lunate, chocolate-brown discal spot and a reddish-
brown or purplish-brown subterminal band. Underside of hind wing concolorous with
forewing, with a slender brown discal lunule and a rather diffuse, reddish-brown
subterminal band; costal margin usually suffused with pink.

Strongly suffused form (stilla Grt.) with normal lines and spots evanescent except
for the prominent median line; area distal to median line heavily overlaid with violet
or reddish-brown.

Expanse: 35.2 + 1.9 mm (50 specimens ).

Egg. Flattened on surface opposite micropyle. Heavily ribbed on micropylar
surface and on sides. Pale greenish-yellow when deposited and remaining essentially
unchanged until a few hours before hatching when larval head gradually becomes
visible through chorion.

Dimensions of egg: height, 0.532 + 0.035 mm; diameter, 0.651 + 0.028 mm (25
eggs).

936 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-6.

Pyrrhia exprimens (Wlk.). 1, 2, Adults, Ottawa, Ont. and Lethbridge,
Alta.; 3, left lateral aspect of ultimate instars; 4-6, dorsal aspect of ultimate instars.

Incubation period: 5 days in all specimens observed.
First-Stadium Larva. Head very dark brown. Prothoracic shield somewhat lighter
brown. Suranal shield medium to dark brown. Trunk cream or light yellow, without

maculation. Setal bases and thoracic legs dark brown; setal bases large and prominent.
\ims of spiracles medium to dark brown.

Head width: 0.355 + 0.013 mm (45 larvae).

Duration of stadium: larvae maturing in five stadia, 6.9 + 1.8 days (21 larvae);
larvae maturing in six stadia, 7.0 + 1.7 days (41 larvae).

cond-Stadium Larva. Head shiny black. Prothoracic shield dark greyish-brown.
| shield greyish-brown. Trunk yellow or cream and usually unmarked; a pair

VoLUME 24, NuMBER 4 OM

of subdorsal whitish-grey lines evident in some specimens. Setal bases, rims of
spiracles and thoracic legs varying from dark greyish-brown to black.

Head width: 0.561 + 0.038 mm (63 larvae).

Duration of stadium: larvae maturing in five stadia, 3.8 + 1.0 days (21 larvae):
larvae maturing in six stadia, 3.7 + 1.5 days (41 larvae).

Third-Stadium Larva. Head completely black, or black marked laterally and in
centre of face with greyish-yellow. Prothoracic shield medium to dark chocolate-
brown with a dull greenish-yellow median line and a pair of lateral greenish-yellow,
submarginal patches; in a few specimens pale submarginal patches so extensively
developed as to almost obliterate brown colouring. Suranal shield medium to dark
chocolate-brown; in some specimens with a median, and a pair of lateral, dull greyish-
yellow spots. Trunk varying from greenish-yellow to rather dark green, with a pair
of pale-yellow dorso-lateral lines; middorsal band often darker than remainder of
trunk. A narrow, pale-yellow, subspiracular line usually evident. Setal bases and rims
of spiracles black. Thoracic legs and proleg shields dark brown to black.

Head width: 0.937 + 0.091 mm (68 larvae).

Duration of stadium: larvae maturing in five stadia, 4.8 + 1.5 days (21 larvae);
larvae maturing in six stadia, 4.9 + 1.4 days (41 larvae).

Antipenultimate-Fourth-Stadium Larva. Head varying from completely black,
through black marked with brown laterally and in centre of face, to black with exten-
sive greenish-fawn areas in middle of parietal lobes. Prothoracic shield greenish-
yellow, typically with four anterior and four posterior black patches; black patches
variably fused. Suranal shield greenish-yellow marked with blackish-brown; least
maculate shield with four anterior and four posterior marginal spots. Middorsal band
green, yellow, or light orange-brown, usually well defined. Subdorsal area consisting
of a median band similar in colour to middorsal band and light yellow marginal lines;
median band of subdorsal area variably suffused with light yellow. Supraspiracular
area concolorous with middorsal band. Spiracular band light yellow, varying greatly in
width and prominence. Suprapodal area concolorous with middorsal band. Midventral
area paler yellow than suprapodal area. Setal bases large and black. Rims of spiracles
dark brown. Thoracic legs and proleg shields dark brown or black.

Head width: 1.31 + 0.07 mm (42 larvae).

Duration of stadium: 3.3 + 1.6 days (41 larvae).

Penultimate-Stadium Larva. Head usually varying from light green to fawn and
marked with black; head occasionally black marked with brown, or less commonly
entirely black. Prothoracic shield pale green to greenish-fawn, marked along anterior
and posterior margins with black; shield occasionally entirely black. Suranal shield
usually poorly distinguished from trunk, occasionally suffused with black. Middorsal
band varying from yellowish-fawn through yellowish-green to bluish-green; usually a
faint, discontinuous, pale-yellow median line through middorsal band. Subdorsal area
yellow with a variably defined, darker, median band; median band similar in colour to
middorsal band, variably suffused with yellow and with a broken and irregular median
line; median band of subdorsal area often so heavily suffused with yellow as to almost
obliterate darker colouring. Supraspiracular area concolorous with middorsal band,
variably and irregularly marked with white or pale yellow. Spiracular band shades of
yellow, usually bright yellow and prominent. Suprapodal area similar in colour to
middorsal band but paler. Midventral area paler than suprapodal area. Setal bases
large and black. Rims of spiracles dark brown. Thoracic legs and proleg shields dark
brown or black.

Head width: 1.81 + 0.17 mm (72 larvae).

Duration of fourth stadium of larvae maturing in five stadia: 4.9 + 1.0 days (21
larvae ).

Duration of fifth stadium of larvae maturing in six stadia: 3.5 + 1.0 days (41
larvae ).

Ultimate-Stadium Larva. (Figs. 3-6). Extremely variable in colouring and macu-
lation. Head varying from pale greyish-yellow, through greenish-yellow, orange-

238 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 7, 8. Pyrrhia exprimens (Wlk.), apical abdominal segments of pupa. 7,
Ventral; 8, right lateral.

yellow and orange to orange-brown; darker heads almost always with black arcs
diverging upward and outward from centre of face and with a black band across frons
and lower surface of genae. Prothoracic shield showing same colour variation as head,
commonly-marked with black spots or patches near anterior and posterior margins.
Suranal shield usually blending with colouring and maculation of trunk. Rims of
spiracles light brown. Setal bases varying from black to concolorous with trunk;
paler setal bases usually each with a dark apical ring. Thoracic legs and proleg shields
varying from black to concolorous with suprapodal area.

Trunk white, yellow, or green variably marked with black; yellow larvae often so
heavily marked with black as to appear predominantly black.

Yellow and black colour phase: Middorsal band black with an irregular and discon-
tinuous, white or yellow, median line. Subdorsal area yellow and white with two
irregular and discontinuous, black median lines. Supraspiracular area dull black
variably and irregularly marked with pale olive-green and white. Spiracular band
white dorsally, pale yellow ventrally. Suprapodal area pale olive-green, variably and
irregularly marked with dull black. Midventral area pale olive-green.

Yellow colour phase: Middorsal band greyish- or brownish-yellow weakly marked
with black. Subdorsal area paler, clear yellow, with a pair of vaguely expressed black
median lines. Supraspiracular area brownish- or greyish-yellow; irregularly marked
with dark brown, particularly along margins. Spiracular band clear light yellow, mar-
gined dorsally with white. Suprapodal area greyish-yellow, lightly and irregularly
marked with clear, light yellow. Midventral area somewhat paler than suprapodal
area.

Green colour phase: Middorsal band apple-green. Subdorsal area creamy-white,
irregularly marked in middle of each segment with pale green. Supraspiracular area
green, irregularly and variably marked with creamy-white. Spiracular band creamy-
white. Suprapodal area green, lightly and irregularly marked with white. Midventral
area greyish-green.

White colour phase: Middorsal band black, discontinuous, often almost completely
suppressed. Subdorsal area dead white with a pair of variably expressed median black
lines; a prominent orange blotch in middle of each segment and extending into mid-

dorsal area. Supraspiracular area black, variably and irregularly mottled with white;

, { - - 1
ick of supraspiracular area often suppressed on at least some segments. Spiracular
ange, often irregularly defined from supraspiracular area. Suprapodal area

VoLUME 24, NuMBER 4 239

dark purplish-brown, variably and irregularly marked with white, usually lightly so.
Midvental area dirty yellow-grey.

Various gradations between these macular forms may also be found.

Head width: 2.86 + 0.11 mm (23 larvae).

Duration of fifth stadium of larvae maturing in five stadia: 7.8 + 1.5 days (21
larvae ).

Duration of sixth stadium of larvae maturing in six stadia: 6.4 + 1.5 days (41
larvae ).

Pupa (Figs. 7, 8). Uniform orange-brown. Spiracles on a level with general sur-
face of cuticle or borne in very shallow depressions. Anterior areas of abdominal
segments 4, 5, 6, and 7 rather heavily pitted; dorsal surface of abdominal segments
1, 2, 3, and 8 moderately pitted. Proboscis terminating between apexes of wings.
Cremaster consisting of two, rather short setae borne at the apex of a conical pro-
longation of the tenth abdominal segment.

Length to posterior margin of fourth abdominal segment: 11.5 + 0.8 mm (14
pupae).

Acknowledgment

I appreciate the assistance of my associate, Mr. E. W. Rockburne in
measuring larval structures, and in drawing the cremaster area of the pupa.

Literature Cited

Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agr. Tech. Bull.
S55 365 pp:

Forses, W. T. M. 1954. Lepidoptera of New York and neighboring states, Part III.
Cornell Univ. Agr. Exp. Sta. Mem. 329, 433 pp.

GARDNER, J. C. M. 1946. On larvae of the Noctuidae (Lepidoptera )—I. Trans. R.
ent. Soc. Lond. 96: 61-72.

Harpwicxk, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6, 116 pp.

PRENTICE, R. M. (compiler) 1962. Forest Lepidoptera of Canada, Vol. 2. Canada
Dept. Forestry Bull. 128, 281 pp.

Watker, F. 1857. List of the specimens of Lepidopterous insects in the collection
of the British Museum, Part 11. London.

A MELANIC ABERRATION OF PAPILIO CRESPHONTES (PAPILIONIDAE )

In 1964, I caught an extreme melanic specimen of Papilio cresphontes Cram. The
specimen was taken in New Orleans during migrating season, probably in November.
It is of normal size and nearly all black with small amounts of white where yellow
should be. The hind wings are nearly all black. The specimen, considerably broken
when caught, was sent to Dr. Alexander B. Klots at the American Museum of Natural
History in February of 1970 for identification. His return letter stated that the museum
has nothing like it, and that although several Society members have recorded melanism,
none are as extreme as this.

VERNON A. Brou Jr., Route 1, Box 74, Edgard, Louisiana.

240 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NEW OR UNUSUAL BUTTERFLY RECORDS FROM FLORIDA

Harry K. CLENCH

Carnegie Museum, Pittsburgh, Penna.

The following records have accumulated in the course of my collecting
in various parts of Florida over the past several years.

Eurema daira daira Godart

Chokoloskee, Collier Co.: 22,31.XII.1967 (14 each, winter form); 19,
20.X1.1969 (4¢é 22, winter form; 24, summer form).

This widespread, common Florida species is of no particular concern
in itself, but these records are of significance in conjunction with the next.

Eurema daira palmira Poey

Chokoloskee, Collier Co.: 22.X1I.1967 (1¢ 12) and 31.XII.1967 (34
32).

All the specimens are of the winter form “ebriola.” This is the West
Indian subspecies of daira and only a few specimens of it have ever been
taken in Florida (Klots 1951; Kimball 1965). These specimens of palmira
raise a question, but unfortunately do not answer it: Is palmira a sub-
species of daira as currently believed, or is it a different, full species?

The possibility that daira and palmira are specifically distinct is sug-
gested by the large number of differences between them (Table 1) and
by the utter absence of intermediates among the specimens of both which
I took in December 1967, all flying in the same area. This possibility is
not diminished by the later (November 1969) captures of d. daira alone
in the same place: there is no evidence whatever of palmira characters
in any of these specimens and hence no evidence of interbreeding in the
intervening time.

The conventional view, that palmira is only a subspecies of daira, is
still possible. The specimens of palmira taken in December 1967 con-
ceivably could all have been offspring of a single immigrant female.
Absence of any evidence of hybridization in later captures could be ex-
plained by swamping, recessiveness of palmira traits, or both.

The present evidence seems to favor their being specifically distinct,
but it is certainly not conclusive. Unless or until we learn that the two

are capable of living together for several generations without loss of
their identities, there is no reason to propose any change in their current
tatus as subspecies.

( omnlic

‘tion should be mentioned. Occasional specimens of d. daira

VoLUME 24, NUMBER 4 241

TABLE 1. Differences between Eurema daira daira and Eurema daira palmira,
based on Floridian and Cuban material. Abbreviations used: up = upperside; un =
underside; fw = forewing; hw = hind wing.

Character

6 2 uphw ground color
é uptw costal ground color

(mixed with gray )

é upfw posterior bar thickness

2 upfw ground color
2 upfw posterior bar

2 uphw terminal fuscous

6 upfw terminal fuscous

& size**

not reaching
Cuz ( thin )
white
absent

thin and absent
below Cui

always extends
beyond Cuz towards
tornus

small

palmira daira
SUMMER FORMS
white yellow*
usually white yellow

broadly reaching
Cuz (thick)

yellow

usually at least

a trace

thick and often
reaches beyond Cup

may extend beyond
Cup, or there be
cut off square

large

WINTER FORMS
1. ¢ 2 uphw ground color white yellow
2. & upfw posterior bar broadly reaching
Cup ( thin ) Cuz ( thick )
always extends cut off square
beyond Cuz towards at Cuz
tornus

not reaching

3. 3 upfw terminal fuscous

4. $9 uphw apical black patch 1 interspace-width 2 interspace-widths
thickness (thin ) (thick )
5. @ upfw ground color white with costal and yellow
apical yellow flush

* Rarely white (see text).

** 10 So of summer daira (Stemper, Fla.): length of fw 16-17 mm, mean 16.5 mm; 10 dc
of summer palmira (Nueva Gerona, Isle of Pines, Cuba): 13.5-15 mm, mean 14.6 mm. Females
of each average larger but show a similar size difference; so do both sexes of the winter forms.

taken in southern Florida (e.g., 24, Toms Harbor, Florida Keys, VIII.
1936 [collector unknown], Carnegie Museum) show white on the upper-
side of the hind wing, a characteristic of palmira. In all other respects
these specimens are true daira and I infer therefore that this is simply
an occasional dimorphic trait, perhaps limited to the warmer areas, and
does not indicate any relationship with palmira.

Urbanus dorantes dorantes Stoll

Chokoloskee, Collier Co.: 20.X1.1969 (26 19, all rather fresh).
This is a new species record for Florida. Kimball (1965) records a
single specimen in the Cleveland Museum which bears two conflicting

242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

labels (Miami, Oct. 11, 1916; and Tampa, June 1908), and is undoubtedly
falsely labelled, as Kimball notes.

Urbanus proteus Linnaeus was ubiquitous and abundant on Choko-
loskee in November 1969, and on the 19th I took a couple “for the record.”
While collecting there again the next day I was belatedly struck by the
fact that some of the specimens seemed to lack the blue-green iridescence
of the dorsal body and wing bases, and took three of these. They turned
out to be dorantes. Although not as common as proteus, they were by
no means rare and additional specimens could easily have been taken.

In view of the range and geographic variation of Urbanus dorantes it
is important to identify the subspecies this Florida population represents
as accurately as possible. Three subspecies are relevant here:

U. d. santiago Lucas (Cuba, Isla de Pinos). Upperside of forewing with hyaline
spots small: bar at cell end reduced to two spots, one or both of which may be
absent; spot below Cus commonly absent. Underside of hind wing with ground color
dark purplish brown (except for paler terminal area), often nearly as dark as the
dark transverse bands.

U. d. cramptoni Comstock (Puerto Rico, Hispaniola). Upperside of forewing with
hyaline spots averaging medium-sized, but varying from as small as in some santiago
to as large as in some d. dorantes: bar at cell end usually entire, rarely divided into
spots and never with one or both spots absent; spot below Cuz occasionally absent,
commonly reduced (much less than half as large as spot in M;-Cu: ), occasionally large
(half or more the size of M:-Cu,; spot); second spot from costa of the subapical three
spots with its distal edge in line with the first or somewhat basad, only rarely a little
distad. Underside of hind wing pale ashy or lavender brown, with dark purple brown
bands, both sexes similar.

U. d. dorantes Stoll (Texas and Arizona south to Paraguay and Argentina). Upper-
side of forewing with hyaline spots large: bar at end of cell always entire; spot below
Cu, always present, large; second spot from costa of the subapical three spots with
its distal edge usually distad of the first, rarely in line. Underside of hind wing pale
ashy or lavender brown with dark purple brown bands; in the female sometimes with
ground much paler ashy.

The three Florida specimens are a small sample but they show these
traits: upperside of forewing with hyaline spots large: bar at end of cell
entire; spot below Cuz present, large; second spot from costa of the sub-
apical three spots with its distal edge distad of the first in all three speci-
mens. Underside of hind wing pale ashy or lavender brown with dark

purple brown bands. The single female has the ground much paler than
the males.
The Florida specimens thus agree in every respect with the mainland
| and South American subspecies, d. dorantes. With Kimball (loc.
cit.) we should certainly have expected any Florida specimens of dorantes
to be of the distinctive Cuban subspecies, santiago. Where, then, did
this population come from? How long has it been on Chokoloskee? How
did it get there? Has dorantes been overlooked elsewhere in Florida (or

—— ene _

VoLUME 24, NUMBER 4 243

the Gulf Coast of Louisiana and Mississippi) because of its resemblance
to the common proteus?

Note. After the manuscript of this paper had been submitted, Dr. Lee D. Miller
wrote that he had captured a female of U. d. dorantes at Homestead, Dade Co.,
Florida, on 19.IJI.1970. This locality is on the opposite side of the state, suggesting

that dorantes is widespread in southern Florida. This, in turn, implies that dorantes
may be a recent arrival or specimens surely would have been taken long before this.

Euphyes dion Edwards

3.1 miles east of Trenton, Gilchrist Co.: 30.IV and 1.V.1968 (5¢ ).

A new species record for Florida. These specimens represent true
dion, not the so-called subspecies alabamae Lindsey, already known from
Florida (Kimball 1965), which I regard as a distinct species following
Clark & Clark (1951). Euphyes dion, however, may need to be racially
divided. Southern specimens are darker below, the ochraceous distinctly
maroon tinged and the fulvous above reduced in extent.

The above specimens were found at Pontederia flowers in a water-filled
roadside ditch, as is discussed more fully under the next species.

Poanes aaroni howardi Skinner

3.1 miles east of Trenton, Gilchrist Co.: 29.IV, 30.IV, 1.V.1968 (134 32).

Records of this species in Kimball (1965) are few and mostly confined
to the autumn. Other localities in Carnegie Museum are: Okeechobee
[Okeechobee Co.], Fla.; Royal Palm State Park [which one?], Fla.; and
a series taken in late June-early July in the northeastern corner of Monroe
Co., Fla.: all leg. W. R. Sweadner.

In view of the temporal dissociation of certain Pennsylvania hesperiines
to avoid competition for flowers (Clench 1967), the association observed
in Gilchrist Co. deserves special comment. The locality lies 3.1 miles east
of Trenton, on Florida Highway 26, and consists of long water-filled
ditches on both sides of the road. The surrounding area is a patchwork
of open marsh, wet meadows (sometimes dry, sometimes inundated),
grassy and sedgy pine-cypress swamp, and dry pinelands. At the time
aaroni and dion were collected the area was in the throes of a drought
that had lasted several months, yet there was still an abundance of water
locally.

I have paid a number of visits to this locality. Three of them were in
late April and early May (29.IV, 30.IV, 1.V.1968). At this time I found
the following Hesperiinae, all fresh, frequenting the roadside ditches and
the Pontederia flowers in them: Atrytone logan Edwards (the common-
est, 19 taken); Poanes aaroni howardi Skinner ( nearly as common, 16
taken); Ancyloxipha numitor Fabricius (6 taken); Ewphyes dion Edwards

944 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(5 taken), Wallengrenia otho Abbot & Smith (3 taken). A few additional
hesperiine species were also taken, but as single specimens only.

Of these, aaroni, logan, and dion in particular were feeding at the blue
flowers of Pontederia, which grew abundantly in the ditches, and on
nothing else. A few other plants were in flower and I repeatedly checked
them, always with negative results.

About five weeks later, on 8 June 1968, I visited the area again. The
Pontederia flowers were almost completely gone and not one of the
hesperiines was found. Two subsequent visits were made in 1969: one
on 19 March, the Pontederia barely emergent above the deep water in
the ditches and not yet in flower; and one on 24 November, when the
ditches were nearly dry, overgrown with grasses, mostly dead, and no
Pontederia flowers at all. Neither of these visits yielded any of the hes-
periines.

These three skippers, logan, aaroni, and dion, may be single brooded
locally, all flying synchronously for a few weeks in the spring, although
the possibility of a second brood in summer or fall in this area is not yet
excluded. The significant point in the present connection is that although
they are competitors these skippers apparently are forced to fly con-
temporaneously by the brief flowering time of their only source of adult
food, Pontederia.

Literature Cited

Criark, A. H. anv L. F. Ciarx. 1951. The butterflies of Virginia. Smithsonian
Misc. Coll. 116 (7), 239 pp.

CiencuH, H. K. 1967. Temporal dissociation and population regulation in certain
hesperiine butterflies. Ecology 48: 1000-1006.

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Gainesville, 363 pp.

Kiots, A. B. 1951. A field guide to the butterflies. Boston, 349 pp.

PIERIS PROTODICE AND URBANUS DORANTES IN
SOUTHERN FLORIDA

Lee D. MILLER AND JACQUELINE Y. MILLER
Allyn Museum of Entomology, 712 Sarasota Bank Building, Sarasota, Florida

Collecting in Florida has uncovered many butterfly and moth species
known from nowhere else in the United States, yet, paradoxically, many
species which logically should be well known from there appear to be

rare or absent. New records from the state are being reported every

VoLUME 24, NuMBER 4 245

year. The implication is clear: Florida’s lepidopteran fauna is still poorly
known, despite all that has been written on it. The observations on the
two species below, taken in Dade and Monroe counties during late March
and early April, 1970, serve to demonstrate this fact rather graphically.

Pieris protodice protodice Boisduval and LeConte

Kimball (1965, p. 37) says about this butterfly, “At present this species
seems to be very rare in Florida.” The majority of his records are from
the northern part of the state, and the species is reported more and
more rarely as one proceeds southward along the peninsula. On the east
coast the most southerly records are from Fort Lauderdale, Broward
County, and Paradise Key, Dade County, where the butterfly is listed
as rare. We found a thriving colony of this species in a grove at the
north end of Homestead, Dade County, and took a number of specimens
from 19 March through 3 April. Because fresh specimens were seen
throughout this period, we feel that the butterfly probably has a long
flight period at Homestead. Had one been concentrating on protodice
it would have taken little time and effort to collect fifty or more speci-
mens in a day, so this butterfly cannot be considered rare there. One of
the foodplants, Shepherd’s Purse, Capsella Bursa-pastoris, a common
weed in the north, was also abundant in the grove, so the checkered
white was not lacking for larval food. Perhaps the recent lowering of
the southern Florida water table has provided a more hospitable en-
vironment for Capsella in disturbed habitats, and protodice may be
spreading southward with the success of its foodplant.

Urbanus dorantes dorantes (Stoll)

Clench (1970) has recorded this species from Chokoloskee, Collier
County, and has listed the first of our specimens from the north end of
Homestead, Dade County. Subsequently we collected eight more speci-
mens from the same area between 30 March and 3 April and saw many
others that were not taken. Two additional specimens were taken along
a trail through a small hammock on Key Largo, 17 miles northeast of
Tavernier, Monroe County, on 31 March. Two others were seen, but
not captured, in the same locality on 4 April. The Key Largo specimens
were taken in the company of such typical hammock species as Eunica
tatila tatilista Kaye and Phocides pigmalion okeechobee Worthington.

Neither Clench (1970) nor we know to which subspecies the purported
Florida dorantes in the Cleveland Museum belongs (Kimball, 1965: 51),
and we doubt the validity of the specimen in any case. We are in full

246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

agreement, however, that the recent specimens are nominate dorantes
(for a discussion of the differences see Clench, 1970), which otherwise
is found from Texas and Arizona southward on the continent, rather than
either of the Antillean subspecies. Just how the continental subspecies,
with an apparent disjunction in range of about a thousand miles by land
and several hundred miles over water, comes to be the Florida subspecies
in preference to the Cuban subspecies, d. santiago (Lucas), with only
a ninety-mile water barrier to cross, is indeed a zoogeographic puzzle.
Postulation of a colonizing flight from Mexico or Texas across the Gulf
of Mexico being successful when no colonization in the same manner
from Cuba has taken place seems beyond the limits of credibility. There
also could be a narrow strip around the Gulf of Mexico from Texas to
Florida in which the species occurs, but if this is true, why have inter-
vening populations not been found? Even in the area from Tampa to
Fort Myers, Florida, where some of the most comprehensive collecting
in the state has been done in recent years by Kimball, King and others,
dorantes has not been detected, so it seems that the idea of interconnected
populations from Florida to Texas is unlikely. The remaining idea, that
dorantes was introduced artificially, seems most reasonable. Certainly
once it got into southern Florida it should thrive, because there is no
shortage of Fabaceae to serve as larval food in the area, and the climate
is comparable with that of other places where this species is native. U.
dorantes must be a relatively recent arrival in the Homestead—Key Largo
area, since it was not recorded by the industrious collecting over many
years by Mrs. Leslie E. Forsyth of nearby Florida City.

Most of the specimens we took were in shaded parts of the two habi-
tats, rather than in bright sunlight, and the butterflies were most com-
monly found during the hottest part of the day. Their habits rather
closely resembled those of the Baja California, Mexico, populations
(Miller, in press) in that they fluttered around the shaded parts of the
area and tended to shun the open places when the sun was brightest.
In such shaded areas dorantes was by far the dominant Urbanus, but in
sunny places U. proteus (Linné) was commonest. The butterflies are
avid flower visitors, preferring blooms of Bidens, so long as these blos-
soms are in comparative shade. Specimens will take up perches and
chase other butterflies in their “territories.” U. dorantes is rather more
wary than proteus, except when they are at flowers.

We would like to thank A. C. Allyn and Miss D. Dunklau for help in
collecting material and H. K. Clench and C. P. Kimball for comments
ind suggestions which led to this paper.

VoLUME 24, NUMBER 4 IAT

Literature Cited

Criencu, H. K. 1970. New or unusual butterfly records from Florida. Jour. Lepid.
Soc. 24: 240-244.

Kmart, C. P. 1965. The Lepidoptera of Florida. Gainesville.

Mitter, L. D. Reports on the Margaret M. Cary-Carnegie Museum expedition to
Baja California, Mexico, 1961. 6. Family Hesperiidae (Lepidoptera). Ann.
Carnegie Mus. (In press).

A NEW SPECIES OF PIRUNA FROM TEXAS ( HESPERIIDAE )

HucH Avery FREEMAN
1605 Lewis Drive, Garland, Texas

During the summer of 1968 J. E. Hafernik collected rather extensively
in the Big Bend area of Texas. Among the many very interesting species
that he collected were two specimens of an undescribed species of Piruna
that is here described.

Piruna haferniki Freeman, new species

MALE (Upper side): Primary dark brownish-black, with seven white, hyaline
spots; a prominent, oval, spot in space 2 and midway between this spot and base of
wing another much smaller spot in same space. In space 3, a small, round, spot
situated directly under apical spot in space 6. Three well developed apical spots
of approximately same size in spaces 6, 7, and 8, forming a straight line; lower spot
directed toward an area between apex and mid termen of wing. A small, upper cell
spot. Fringe light brown, uncheckered.

Secondary dark brownish-black, unmarked. Fringe light brown, uncheckered.

MALE (Under side): Primary brown, slightly lighter in the apical region. White,
hyaline spots present and better defined than on upper side.

Secondary dark brown with a heavy overscaling of purplish-gray scales over basal
half of wing and extending from anal fold along outer margin to space 5. A slightly
ferruginous area extending above space 5 along outer margin to apex. Throughout
discal area from space 1 to costa an area without overscaling; here brown ground
color forming a broadly triangular area darker than the remainder of the wing. No
spots present.

BODY: Thorax dark brownish-black above, lighter brown beneath. Abdomen dark
brownish-black above, lighter brown beneath. Head brown, with a few yellowish,
hair scales present. Palpi white at base, yellowish at termen, dark grayish on lateral
sides. Legs golden brown. Antennae, both shaft and club, dark brownish-black
above, lighter beneath; club especially pale beneath, yellowish. Apiculus golden.

Wing measurements. Primaries: base to apex, 11 mm; apex to outer angle, 7.5 mm;
outer angle to base, 9 mm. Secondaries: base to end of vein 3, 9.5 mm; center of
costa to anal angle, 7.5 mm. Total expanse: 23 mm.

Type Material: Holotype, male, Green Gulch, Big Bend National Park, Brewster
County, Texas, elev. 5700 ft., 4 August 1968 (J. E. Hafernik) in the American
Museum of Natural History, New York. One male paratype, same location and col-
lector, 24 July 1968, at present in my collection.

948 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

dL ry

Figs. 1 and 2. Upper and under side of Piruna haferniki Freeman, holotype, male,
Green Gulch, Big Bend National Park, Brewster County, Texas, elev. 5700 ft., 4
August 1968 (J. E. Hafernik).

Superficially on the upper side P. haferniki resembles P. cyclosticta
(Dyar); however the maculation is much better developed in haferniki.
There are two basic features in haferniki which readily separate it from
any other Piruna: (1) In all other species of Pirwna, the apical spot in
space 7 is either smaller than the other two or else absent, whereas in
haferniki it is approximately the same size as the others; in all other
Piruna the line formed by the apical spots is directed toward the center
of the outer margin of the wing instead of between this area and the
apex as it is in haferniki. (2) No other Piruna has the unusual purplish-
gray overscaling on the under side of the secondaries leaving the darker
brown discal area which is present in haferniki.

There are basic differences in the genitalia, especially the uncus, sepa-
rating haferniki from the other known species of Piruna. P. haferniki
is a member of the brunnea complex which contains brunnea (Scud.),
cyclosticta (Dyar), and pirus (Edw.), and can be readily distinguished
from these three species by the above characteristics.

Fig, 3. Piruna haferniki, male genitalia of paratype, Green Gulch, 24 July 1968
( Hafernik ).

VOLUME 24, NuMBER 4 249

Acknowledgments

The author wishes to thank the American Philosophical Society for a
grant which is making research on the Hesperiidae possible. The photo-
graphs of the adults used in this article were made by Mr. William L.
Skinner, Physics Teacher, Hillcrest High School, Dallas, Texas.

Literature Cited

Draupt, M. 1924. Hesperiidae. In Macrolepidoptera of the World. Ed. by A.
Seitz, Vol. 7. Stuttgart.

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the clas-
sification and nomenclature adopted in the British Museum. Part 4. Hesperiinae
and Megathyminae. London.

GopMan, F. D. Anp O. Satyin. 1887-1901. Biologia Centrali-Americana. Insecta.
Lepidoptera—Rhopalocera 2: 244-637.

OBSERVATIONS ON SOME PHYCITINAE (PYRALIDAE) OF
TEXAS WITH DESCRIPTIONS OF TWO NEW SPECIES

ANDRE BLANCHARD
P.O. Box 20304, Houston, Texas

The latest revision of the Phycitinae is that of Heinrich (1956). It in-
cludes a description and figures of genitalia of every New World species
known at the time of its publication. It makes it possible, for anyone
who can dissect genitalia, to identify a very large proportion of his cap-
tures and to call for expert advice in difficult cases only.

In trying to classify my Phycitinae I have been puzzled by several
species which do not appear to be included in Heinrich’s monograph.
For all of these, except two, I have only one specimen or specimens of
only one sex. These will have to wait a little longer. This paper offers
a description of the two new species of which I have males and females;
it also includes an annotated list of some species which Heinrich had
not seen from Texas or even from the United States.

Zamagiria kendalli A. Blanchard, new species

Male: Tongue well developed. Antennae brown, shortly ciliate, a deep sinus at
base of shaft, fringed with heavy scale tufts, bearing on each of the three upper
segments a claw-like, well-sclerotized process (Fig. 3). Head slightly depressed be-
tween antennae and in upper part of front. Labial palpi recurved ascending, first
segment whitish, second segment sprinkled with brown scales, very long, broadly
dilated and hollowed within to hold maxillary palpi, third segment minute (Fig. 3a).
Maxillary palpi in the form of large aigrettes. Head, collar and disc of thorax dark

50

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

VOLUME 24, NuMBER 4 Zon

Fig. 7. Zamagiria kendalli, holotype.

Fig. 8. Dasypyga salmocolor, holotype.

<

Figs. 1 to 4. Zamagiria kendalli. 1, Male genitalia, aedeagus omitted; la, aedeagus;
lb, transtilla; 1c, tufts of eighth abdominal segment; 2, female genitalia; 3, part of
male antenna; 3a, denuded male labial palp; 3b, denuded female palp; 4, venation
(female wings ).

Figs. 5 to 6. Dasypyga salmocolor. 5a, Male genitalia moderately expanded,
aedeagus omitted; 5b, male genitalia fully expanded, aedeagus omitted; 5c, aedeagus;
6, female genitalia.

(Scale shown on all figures is one millimeter except for 1c and 4.)

952 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

brownish gray; patagiae darker; abdomen above lighter. Underside of thorax, ab-
domen and legs grayish white. Primary (Fig. 7) with ground color pale bluish gray,
heavily dusted, mostly between radial vein and inner margin, with brownish to red-
dish and a few black scales. T.a. line well defined only between radial and anal
veins, bluish white; narrowly bordered distally by a black line bending basad, almost
squarely, over the radial and running to costa; narrowly bordered basally by a black
line meeting a broader, black basal dash on anal vein. S.t. line bluish white, best
defined by its black inner border deeply and sharply indented opposite upper angle
of cell and on Cu2; this last cusp bringing it below discocellular vein. A crescent
shaped spot, with ground colored center, entirely outlined in black at the end of
cell, the more basad arc of the crescent contour lying directly over discocellular vein.
Black, intervenular, almost confluent, terminal dots. Fringe light brownish gray.
Secondary grayish white, slightly darker at apex and along termen. Fringe light gray;
the shorter scales, being dark tipped, produce a median blackish line. Undersurface
of primary almost uniformly brown, basal half of costa darker. Secondary iridescent
grayish white, brown between costa and radial vein and near apex.

Female: Similar to male except for the following: antennae simple pubescent,
labial palpi (Fig. 3b) not hollowed out, ascending but not exactly applied against
front, maxillary palpi minute, filiform.

Venation: As shown in Fig. 4.

Expanse: 20 to 24 millimeters.

Male genitalia: As represented by Fig. 1, la, lb. Uncus broadly triangulate.
Gnathos elongate, arising well down from ventrolateral projections of tegumen, ter-
minating in a spatulate, hollow hook. Transtilla absent. Aedeagus stout. Penis
armed with one or two strong cornuti. Eighth abdominal segment with ventral tuft
as shown in Fig. lc.

Female genitalia: As shown in Fig. 2.

Of all the genera included in Heinrich’s monograph, Zamagiria appears
to accommodate kendalli best: the venation, labial and maxillary palpi,
male antennae, as well as the male and female genitalia indicate a close
relationship. The head, however, in the upper part of the front and be-
tween the antennae, is somewhat depressed rather than deeply grooved,
and the “appressed, matted tuft of long scales arising from the upper
edge of the frons” is missing. Although the male genitalia differ markedly
from all those figured by Heinrich (which, as he remarks, exhibit striking,
structural, specific differences among themselves) they agree quite well
in general characters. The combination of elaborate wing masculation
and genitalia differentiate this species very definitely from all previously
described taxa.

Food plant: On 7 July 1969 in Musquiz Canyon (Tex. Hwy 118)
N.W. of Alpine, Texas, Roy and Connie Kendall found several larvae,
sheltered inside folded or rolled portions of leaves of Croton fruticulosus
(Engelm.). They reared three with the following results: pupated 10
July (1), 2 emerged 23 July; pupated 12 July (1), ¢ emerged 23 July;
pupated 18 July (1), 2 emerged 28 July.

[ have these three specimens before me; as they are definitely con-
specific with those taken in our light traps, they are labeled paratypes.

VOLUME 24, NuMBER 4 253

It gives me pleasure to name the new species for our dear friends, who
have discovered the life histories of many Texas species of Lepidoptera,
and with whom we have enjoyed many fruitful and pleasant field trips.

Type material: Holotype male, Fort Davis, Hospital Canyon, behind historical
fort (Jeff. Davis Co.), Texas, 5000’, 11 July 1969 (genitalia on slide A.B. 2050)
deposited in the U.S. National Museum (No. 71004).

Eighteen paratypes, including the three listed above in the Kendall collection:
Fort Davis, 16, 39 9, 24 Aug. 1967; 16, 19, 11 Jume 1969; 36 6,59 9, 11 July
1969; Guadalupe Mts., Bear Canyon, 5700’, Culberson Co., Texas, 13, 4 Sept. 1969.

.
>

Dasypyga salmocolor A. Blanchard, new species

Head covered with ochreous salmon scales, paler and thinner on front than on
vertex, with a transverse parting line behind antennae. Tongue well developed. An-
tenna weakly pubescent. Labial palpus upcurved, not quite reaching vertex, very
slightly longer in female than in male; third segment a trifle longer than second,
acuminate. Maxillary palpus small, squamous. Thorax with collar, disc of thorax,
and patagiae concolorous with vertex. Legs closely scaled, varied with bright ochre-
ous salmon, purplish black, and whitish. Abdomen pale ochreous salmon; with some
blackish scales on the more basad segments beneath. Upper surface of primary (Fig.
8) ochreous salmon. T.a. line pale purplish gray, starting on costa one fifth distance
from base to apex, nearly straight to inner margin, and reaching latter one fourth
distance from base to anal angle; rather wide, and narrowly bordered on both sides
by sprinkling of darker scales. A small patch of black, coarse, raised scales in sub-
median fold on inner side of t.a. line. Basal area soiled in places with purplish scales,
some of them slightly raised. On male specimens, ground color somewhat darker
along outer border of t.a. line, on a width about equal to width of t.a. line itself.
S.t. line absent, but along termen a border about same width and color as t.a. line,
continuing in a rather wide, concolorous fringe. No discal dots. In cell M: a fine
red line running nearly to outer dark border, and thence angling sharply to apex. A
similar red line on lower fold in outer area. Secondary pale, shining, smoky fuscous;
somewhat darker along outer margin and toward apex. Fringe very pale smoky fus-
cous, with a fine dark line consisting of the tips of the shortest scales. Undersurface
of primary almost uniform ochreous. Secondary almost uniform paler ochreous.

Expanse: 19-20 mm.

Male genitalia: As represented by Figs. 5a, 5b, 5c. Fig. 5a showing those of the
type prepared with moderate expansion; Fig. 5b showing those of another specimen
fully expanded to reveal inner face of valve; 5c showing aedeagus of same specimen.

Female genitalia: As represented by Fig. 6.

Type material: Holotype male, Sierra Diablo Wildlife Management Area, 6000’,
Culberson Co., Texas, 1 September 1969 (genitalia on slide A.B. 2079); deposited
in the U.S. National Museum (No. 71005).

Five paratypes, all from same locality: one male, 6 June 1969; two males 8 June
1969; one male 14 July 1969; one female 1 September 1969 (genitalia on slide A.B.
2080).

Dasypyga salmocolor is extremely close to D. alternosquamella, Rago-
not. It can be distinguished from the latter by its much reduced dark
basal area, extending definitely less than a fourth, instead of almost one
half, the length of the wing. The male genitalia also show the close rela-
tion between the two species, but the shape of the free arm of the sac-
culus is quite different.

254 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Other Interesting Records

I will now list some Phycitinae which Heinrich did not see from Texas
or in some cases even from the United States, which my wife and I have
collected in our effort to prepare a catalogue of the moths of Texas.

Acrobasis kearfottella Dyar: Tennessee Colony (Anderson Co.), Conroe
(Montgomery Co.), Huntsville State Park (Walker Co.), end of May.
These records extend the distribution indicated by Heinrich more than
five hundred miles.

Acrobasis stigmella Dyar: Tennessee Colony (Anderson Co.) end of
May. An extension of range of over five hundred miles.

Bertelia grisella Barnes & McDunnough: My identification of this in-
sect has been delayed a long time by what I think is a poor representa-
tion of the transtilla in Heinrich’s Fig. 187. I have a series caught at
Shafter (Presidio Co.) and a few specimens from the Guadalupe Mts.
(Culberson Co.), all taken in October.

Myelopsis alatella (Hulst): Seen only once, but was abundant, at
Belton Reservoir (Bell Co.), 6 April 1970. Nearest locality indicated by
Heinrich: Pecos, New Mexico.

Fundella ignobilis Heinrich: I have only one female specimen. The
female genitalia leave no doubt about its identity. It was caught five
miles southeast of Mt. Livermore in Jeff Davis Co. This is a new record
for the U.S. It was previously taken in Central America.

Anadelosemia condigna Heinrich: Shafter (Presidio Co.), Big Bend
National Park, Davis Mts. (Jeff. Davis Co.), October. Described from a
dozen specimens caught in the Baboquivari Mts. of Arizona.

Zamagiria australella (Hulst): Two specimens only, from Santa Ana
National Wildlife Refuge and from the Welder Wildlife Foundation, both
taken in mid-November. The type in the U.S. Nat. Mus. is labeled
Blanco, Texas but Heinrich suspected that it might be a Florida specimen.
There does not seem to be any reason for this doubt any longer.

Ancylostomia stercorea (Zeller): Brownsville (Cameron Co.), mid-
October, two males.

Pima albiplagiatella occidentalis Heinrich: Fort Davis (Jeff. Davis
Co.), end of March, two males, one female.

Pima granitella (Ragonot): Fort Davis (Jeff. Davis Co.). Big Bend
National Park (Brewster Co.), Mt. Locke in Davis Mts., end of March;
Paducah (Cottle Co.) mid-April.

Quasisalebria admixta Heinrich: This is quite common in the Sierra
Diablo Wildlife Management Area and in the Guadalupe Mts. (both in
Culberson Co.) in June and September. Only one female was taken in
Big Bend Nat. Park in October.

VoLUME 24, NuMBER 4 955

Actrix nyssaecolella (Dyar): Tennessee Colony (Anderson Co.) mid-
April, one female.

Actrix dissimulatrix Heinrich: Huntsville State Park (Walker Co.),
May, one female.

Dioryctria auranticella (Grote): Three specimens from near the top of
Mt. Locke in the Davis Mts. early July.

Patriciola semicana Heinrich: Sierra Diablo Wildlife Management
Area (Culberson Co.) end of September; Davis Mts. (Jeff. Davis Co. )
early October.

Anderida sonorella (Ragonot) (?): Sierra Diablo Wildlife Manage-
ment Area (Culberson Co.) early June and late September; Mt. Locke,
Davis Mts. (Jeff. Davis Co.) September and October; Paducah (Cottle
Co.) July. All my specimens have a sprinkling of rusty red scales along
the inner margin of the primaries, which is not mentioned by Heinrich;
yet their genitalia, male and female, agree with his figures.

Homoeosoma impressale Hulst: Fort Davis (Jeff. Davis Co.) May.
Paducah (Cottle Co.) April.

Unadilla erronella (Zeller): Brownsville (Cameron Co.) November.
Big Bend Nat. Park ( Brewster Co.) October. A surprising record as Hein-
rich had it only from Central America and the Greater Antilles. Yet the
female genitalia leave little doubt about the correctness of this identifi-
cation.

Rhagea packardella (Ragonot): Not uncommon from June until Sep-
tember in my collecting places of West Texas.

Eremberga insignis Heinrich: Not uncommon at Big Bend Nat. Park
in September and October but does not seem to reach even Fort Davis.
A new U.S. record.

Acknowledgment

It is a pleasure to acknowledge with warm thanks the help given me by
Dr. D. C. Ferguson of the Systematic Entomology Laboratory, U.S.D.A.
in examining some of my specimens and revising part of this paper.

Literature Cited

Heinricu, Cart. 1956. American Moths of the subfamily Phycitinae. U.S. Nat.
Mus. Bull. 207.

256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

INCISALIA FOTIS SCHRYVERI (LYCAENIDAE): BIONOMIC
NOTES AND LIFE HISTORY

CuirForD D. FERRIS
University of Wyoming, Laramie
and

Ray E. STANFORD

Denver, Colorado

In the years that have elapsed since the original description of Incisalia
fotis schryveri Cross (1937), little additional information has been gath-
ered regarding the biology of the insect. The distributional limits have
been imprecisely determined, the immature stages have remained un-
described, and only vague speculation has appeared regarding possible
host plants. This paper constitutes the first description of the immature
stages and a record of the host plant. The insect was studied in Wyoming
(by Ferris) and in Colorado (by Stanford). In the paragraphs which
follow, where regional differences exist, state names will be mentioned;
otherwise descriptions pertain to the entire range of schryveri.

Ecology and Nature of Habitat

Incisalia fotis schryveri occurs in multiple colonies in the eastern foot-
hills of the Front Range of the Rocky Mountains in north-central Colo-
rado, and in the continuation of this range into south-eastern Wyoming.
Its northern limits, or possible blend zones with I. fotis mossii (H.
Edwards), have yet to be determined, but in Colorado it seems to ex-
tend no farther south than El] Paso Co. Records are also available from
Boulder, Clear Creek, Douglas, Gilpin, and Larimer Cos. The species
probably occurs also in the northeast portion of Park Co., along the Platte
River, and may be found in parts of Teller Co. In Wyoming, it is known
from Albany, Carbon, and Converse Cos., and is associated with the
Laramie, North Platte, and Platte River drainages. Figure 1 shows records
as dots and suggested distribution as shading.

The habitat of I. fotis schryveri in Colorado has been characterized
briefly by Eff (in Brown, 1957). The species occurs in the Transition
Zone, being found in sparsely wooded, brushy foothill canyons ranging
in c.ovation from about 5,600 to slightly over 8,000 feet. The terrain con-
sists it neral of east-facing canyons containing intermittent or perma-
nent strea eparated by steeply sloping rocky ridges. The predominant
“round cover is Cercocarpus montanus Raf.; other characteristic vegeta-
uniperus scopulorum Sarg. (south slopes), Pseudotsuga

VoLUME 24, NuMBER 4 PART)

CARBON

MQUSS SSO}

Vv
SLA S

YG SIBLH VS AG GAAS

WYOMING

COLORADO

44 FASO

Fig. 1. Known collection areas (dots) and probable distribution (shaded areas)
of I. fotis schryveri. Wyoming: Albany Co.: Pole Mtn. E. of Laramie; Carbon Co.:
20 miles S.E. of Encampment; Converse Co.: Natural Bridge. Colorado: Boulder
Co.: Flagstaff Mtn., Boulder Canyon, Four-Mile Canyon, Gregory Canyon, Mesa
Trail, Sunshine Canyon, Two-Mile Canyon, Sugarloaf, North St. Vrain Canyon; Clear
Creek Co.: Clear Creek Canyon; Douglas Co.: Jarre Canyon nr. Sedalia, Platte Canyon;
El Paso Co.: nr. Palmer Lake, N. Cheyenne Canyon; Gilpin Co.: Blackhawk Creek;
Jefferson Co.: Chimney Gulch (type locality, large dot on map), Coal Creek, Golden,
Lookout Mtn., Platte Canyon, Bear Creek Canyon, West of Morrison, Red Rocks Park,
Clear Creek Canyon, Golden Gate Canyon, Indian Gulch, $.W. of Morrison, S. Platte
Canyon nr. Waterton.

258 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

menziesii Franco (north slopes), Pinus ponderosa Laws., Celtis occi-
dentalis L., Mahonia repens G. Don, Ceanothus fendleri Gray, Ribes,
Rhus, Yucca, Opuntia, Eriogonum umbellatum Torrey, and Chrysotham-
nus. Typical riparian shrubs and trees include Acer negundo L., Prunus,
Crataegus, and Salix. Schryveri seldom strays more than a few hundred
yards out into the prairie, is distinctly uncommon in habitats dominated
by Quercus gambelii Nuttall, and is nearly totally replaced by three other
elfins (I. polios Cook and Watson, I. iroides (Boisduval), and I. eryphon
(Boisduval) ) above 8,000 feet. Of special interest is the occurrence of
the stonecrop, Sedum lanceolatum Torrey, in the habitat, as it serves as
the host plant for schryveri. Although widespread at higher elevations
in the Rocky Mountains (where it serves as host plant for Parnassius
phoebus sayii Edwards ), stonecrop occurs in small, rather isolated clusters
in the foothills. These are found mostly on well-drained east or northeast
slopes with slight water seepage, and tend to be associated closely with
M. repens, Orobanche ludoviciana Gray, and E. umbellatum. According
to Eff (in litt.), the last of these associated plants may serve as host for
Callophrys sheridanii (Edwards), a species sympatric and synchronic
with schryveri in the region discussed in this paper.

Schryveri and sheridanii are usually the first non-hibernating species
to appear in the spring, often in late March. (Artificial eclosion was ef-
fected by Stanford on 3 February 1970 by bringing pupa into a warm
room.) The flight period is about three weeks, but stragglers may be
found in late May or even early June. A fresh male taken by Stanford
in Clear Creek Canyon, Jefferson Co., on 7 September 1968 is considered
an atypical emergent rather than evidence of a second brood. The April
flight period in Colorado corresponds with the blooming of B. repens, O.
ludoviciana, and Salix species, but schryveri is seldom seen at flowers.
Males tend to congregate in canyon bottoms, where they perch on various
twigs or upon the ground. They exhibit markedly pugnacious behavior,
chasing any other butterflies in the vicinity, and even an occasional bird.
Frequently a specimen of Nymphalis antiopa (L.) or of Polygonia
zephyrus (Edwards), both common species in the habitat, will be as-
saulted by dozens of schryveri simultaneously. Occasional females are
seen in the canyon bottoms, nearly always fresh and either virgin or in
copulo, but to find them in numbers one must ascend the steep hillsides
to the slopes where Sedum lanceolatum occurs. Here the females are

abundant, especially in the early afternoon. They fly close to the ground
and alight frequently upon stones or plants of many kinds. Oviposition
has been observed only upon Sedum, where a female will deposit a single

egg upon the superior surface of a basal leaf. Of nearly fifty eggs re-

VoLUME 24, NumBer 4 259

Fig. 2. a, Wyoming habitat. b, Colorado habitat. c, Host plant (in situ).

covered in various foothill habitats west of Denver in 1969, all but one
or two were so situated. A few Sedum plants were found bearing two
or three eggs, and an occasional egg was found on the ground beneath
or nearby a Sedum plant. Maximum concentration of eggs was observed
during the last week in April, and by May 10th, only larvae were found
on the plants. The early instars fed upon the leaves, both in the field
and in the laboratory, but by mid-May, when the plants were in early
bloom, the by-then-third or fourth instar larvae preferred the flowers and

260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

buds to the leaves. The pink-buff color of the third and fourth instar
larvae is nearly identical to the color of the unopened buds of the Colo-
rado morph of S. lanceolatum, while the pale green of the early instars
matches the basal leaves, an interesting example of the changing of pro-
tective coloration. (See the description of the larvae in Wyoming which
follows for comparison. )

In Wyoming, this little butterfly inhabits well-drained slopes and hill
crests in the Upper Transition-Lower Canadian Zones. The vegetation
characteristic of the region is sagebrush (Artemesia species), various
grasses, Pinus flexilis James, Pinus ponderosa Laws., Juniperus scopulorum
Sarg., Populus tremuloides Michaux, Arctostaphylos uva-ursi L., Sedum
lanceolatum Torrey, and various annual and perennial herbs. The eleva-
tion ranges from 8,000 to 8,500 feet in most of the habitats.

I. fotis schryveri makes its appearance along with C. sheridanii while
there are still patches of snow on the ground. It is heralded by the flow-
ering of the sagebrush buttercup (Ranunculus glaberrimus Hook.) and
pasque flowers (O. ludoviciana also known as Anemone patens L.). In
1969, schryveri was first collected on May 4th with four females and six
males being taken. The last specimen, a female, was taken on May 23rd.
No examples of this species were seen after this date. In Wyoming, it
is sympatric with I. polios and I. eryphon, although these elfins appear
on the wing about midway in schryveri’s flight period.

The butterfly is invariably found on the slopes with eastern or south-
eastern exposure. It appears to be somewhat more sensitive to weather
conditions than other species with which it is sympatric. Depending upon
air temperature, flight begins as early as 9:00 A.M. (MST) and extends
into the afternoon until the first shadows fall over its habitat. It dis-
appears immediately if the sun becomes obscured by clouds. The males
are fond of flying about and resting upon low sagebrush plants. They
may also be found sunning themselves (wings folded over back) on sandy
patches of soil. The females are less active and keep quite low to the
ground, frequently resting upon the ground or the host plant. They are
most active about noon when they may be seen ovipositing. As in Colo-
rado, the males are prone to combat and may be seen fighting with one
another, or with other species, over the tops of the sagebrush clumps.
The flight of schryveri is quite erratic, although never more than a few
teet above the ground. Males and females are found in the same loca-
tion in about equal numbers (cf. Colorado). The species is quite local
in Wyoming and the presence of the foodplant does not indicate that the
insect will be found. When located, both the insect and the foodplant

il! generally be in high concentration.

VOLUME 24, NUMBER 4 2961

d

Fig. 3. a, Ovum on seedum leaf. b, Second instar larva prior to moulting. c, New
third instar larva. d, Pupa, ventral aspect. e, Fourth instar larva in characteristic
head down feeding position; note intensity of chevron markings. f, Adult male, dorsal
and ventral surfaces. g, Adult female, dorsal and ventral surfaces.

262 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY

Ly
4%,
&

5

4%

y
5

5
%

soho?

a EEE

A

Fig. 4. a, Ovum. b, First instar, dorsal view and transverse section through a
middle segment. c, Second instar, dorsal view, transverse section through a middle
segment, angled view of dorsal surface of three segments showing tubercles. d, Third
instar, lateral and anterior views. e, Fourth instar, ventral aspect just before pupation.
[, Pupa, dorsal aspect. g, Pupa, ventral aspect. h, Pupa, lateral aspect.

Mii

The foodplant grows quite close to the ground, frequently under the
clumps of sagebrush. In Wyoming, S. lanceolatum is distinctly poly-
morphic. At low elevations (7,500’), the plant is dark green with deep
rose leaf tips and crown. In the Laramie Mountains in the schryveri

VoLUME 24, NuMBER 4 263

habitat, stonecrop is a yellowish gray-green with the leaf tips only faintly
rose tinted and the crown yellow-green. Plants from the Sierra Madre
Mountains (Carbon Co.) are distinctly rose to rust colored with yellow
crowns. The flowers vary from pale yellow-green to yellow in the Laramie
Mountains, and are bright yellow in the Sierra Madre Mountains.

The differences in larval feeding and adult flight patterns between the
Colorado and Wyoming insects would appear to be simpiv adaptations
to different habitats as determined by the location and morphology of
the host plant.

Life Study

The description of the life stages which follows is based upon material
collected in 1969 on the slopes of Pole Mountain, Sherman Range of the
Laramie Mountains in Albany County, to the east of Laramie, Wyoming.
This is an area in which schryveri has not been previously reported. The
study is based upon 37 males and 32 females taken in the field. In all
respects, excepting feeding habits of the larvae, as discussed previously
and below, the immature stages of the Colorado and Wyoming insects
are similar.

On 16 May, one egg was located on the foodplant, S. lanceolatum. On
17 May, a female was captured while ovipositing. On that day and the
following day, three additional living females were taken. Additional
eggs were secured from the habitat as well. The four females all proved
to have been mated although two appeared freshly emerged. In all, about
100 eggs were obtained. The insects showed no reluctance to ovipositing
in captivity. They were kept in widemouth specimen jars (screened
tops) into which soil and the foodplant had been introduced.

Ovum: The eggs are oblate spheroids, pale green, about 0.8 mm in diameter.
They are deposited in the natural state singly on the larger clusters with only one
egg per Sedum cluster, which makes their location in the field somewhat tedious and
time consuming. In captivity the females deposited on a single plant cluster until
exhausted. In Wyoming, the one-egg-per-plant cluster in the wild state seems to be
a natural protection. The larvae are extremely voracious and the foodplants are
quite small in early spring. It was observed that one larva could, during its four
instars, devour a single cluster of the host plant. Since, in the field, the clusters of
Sedum may be separated by several feet, the one egg per cluster rule seems to be
a guarantee of a food source to the developing larva. The eggs (in the wild) are
normally deposited at the base of a leaf on the under surface (cf. Colorado observa-
tions). Eggs were deposited in captivity from 18 to 22 May, 1969. The eggs were
maintained at an average temperature of 21° C with a plus or minus 5° C (approxi-
mately) daily variation. In the field at this time, the temperature range was from
about 2° C to 24° C (air temperature) for the habitat. Twenty-four hours prior to
emergence of the larvae, the eggs turn in color from pale translucent green to opaque
white.

First Instar: The first larva, from the eggs of 17 May, was observed on the morn-
ing of 23 May. The first instar is illustrated in Figure 4. Initially the larva is about

264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1 mm long and pale yellow-green. Within 24 hours, the larva has reached 2-3 mm
and the color has changed to pale tan or dark buff. The hairs are uniformly black.

Second Instar: Initially growth is quite rapid and the first larval moult of the
23 May larvae occurred on 25 May. The second instar is rose-pink and white. The
pink approximates the color of the leaf buds at the top of the foodplant. Initially the
larvae spend most of their time at the tops of the Sedum clusters. The hairs are now
neutral light color. In this instar, larval length increased from 3 to 5 mm.

Third Instar: Larvae hatching on 23 May moulted into the third instar on 27
May. The basic colors were similar to those of the second instar but with a pattern
change and more pink than white. The larvae have a very liquid appearance at this
point. The color subsequently changes into a medium rose-pink. The light portions
dorsally are very light cream while ventrally the color is a medium cream. During
this instar, the length increased from 5 mm to 1 cm.

Fourth Instar: The third and fourth instars are very similar, and for this reason
it was difficult to note exactly when the moult between them first occurred. It ap-
peared to be 6 June. The last of the larvae made the transition on 13 June. The
colors are somewhat muted relative to the third instar. The light areas are somewhat
darker and there is an overall undertone of greenish. Just prior to pupation, the
ventral areas become a dark cream color. In both the third and fourth instars, the
hairs are neutral (straw) color. The larval size increased from 0.8 mm (minimum
value of maximum length of third instar) to 2 cm. Prior to pupation, the larvae spin
a fine light mat of silk and shrink in size to 1 cm. They assume the ovoid shape of
the pupa. The prepupal stage lasts from 24 to 72 hours. During the premoult period
of the last two larval stages, the caterpillars also spun light mats and remained in-
active for period up to 24 hours.

Pupa: The first pupa was formed on 13 June, thus giving the period from deposi-
tion of egg to pupation as 28 days. The average length of the pupae is 0.9 cm.
Initially the wing areas of the pupa are medium cream in color and the abdominal
area is bright pink. Within 24 hours, the color has changed to a uniform medium
brown. In captivity, the pupae were formed on the glass sides of the rearing jars, on
the soil in the open, or under debris. They were loosely attached to light silk mats.
The larvae exhibited a gregarious nature before pupation and clusters of from 6 to 8
pupae were common. A total of 84 pupae were obtained, the last on 4 July.

The various stages of development are illustrated in the accompanying
figures.

In Colorado (material collected and observations made in April-May,
1969, Chimney Gulch (type locality) and Indian Gulch, Jefferson Co. )
pupation occurred between 27 May and 8 June in the laboratory; the
average duration of the egg stage was 6 days (5-8), and of the larval
stage 31 days (27-35). Field observations were too few to establish the
length of the larval stage in nature, but the fact that fourth instar larvae
were found as late as 14 June would suggest a slightly longer maturation
time in the cooler natural environment than in the 22—24° C laboratory.
Pupation occurs on the ground, often several feet away from the host
plant which served a given larva, and usually in a semi-sheltered spot
such as near to a small stone. No pupal dissections were made to deter-
mine the stage in development when diapause takes place, but the fact
that reared insects emerged from pupae within 2-4 days of their transfer
from outdoors to indoors (Stanford, Denver, Colo., Feb., 1970) suggests

VoLUME 24, NuMBER 4 265

that imaginal development is nearly complete at diapause. Some of the
Wyoming pupae showed the wing pattern of the imago within a week
after pupation.

Adult: The mature insect is illustrated (Fig. 3:f,g). It is here de-
scribed in detail because Cross (1937) failed to do so in his original
description:

Dorsal primaries of male uniformly warm gray-brown with pad just slightly paler
than ground color; dorsal primaries of female generally fulvous with apex, costal and
outer margins warm gray-brown; dark color of apical area extending into subapical
region. Both sexes with fringe white with some black hairs at the vein ends. Dorsal
secondaries of male with basal and discal areas uniformly warm gray-brown; lower
half of limbal area, extending over anal angle, and approximately one-third of lower
anal margin dark fulvous; female generally fulvous over entire dorsal surface with
some darker scaling in basal area and along anal margin. Fringes in both sexes same
as in primaries.

Ventrally sexes similar. Primaries with apical area extending along outer margin
a rich brown tending to gray-brown at the tornus. Limbally, black oval spots in cells
Cu, to Rs as a submarginal row. Postmedian area brown suffused with gray and white,
and separated from the median portion by an irregular band which is white distally
and almost black radially (basally). Discal area gray-brown tending to brown, with
dark scales in basal area and with fulvous scales toward the irregular band. Secondaries
with irregular band repeated. Discal and basal areas a warm brown with some gray
suffusion. A submedian dark irregular band. Postmedian area gray-brown heavily
suffused with white. A submarginal row of black spots, diminishing in size, one in
each of cells to Cus. Remainder of limbal area brown irrorate with lighter scales
basally and darker scales distally. A thin white band along the outer margin followed
distally by a brown band showing enlargement along vein ends. Fringes as in dorsal
aspect.

Head, crown, and front with dark gray-brown scaling in male and red-brown scaling
in female; eyes black, hairy, bordered (ringed) with white. Thorax and abdomen
concolorous with dorsal wing surfaces, and hairy. Underside of abdomen heavily over-
laid with white hairs. Legs, dark gray-brown shot with white.

Wingspan of specimens in authors’ collection: males and females (LFW) 1.3-1.4
cm.

The specimens from Colorado are not significantly different from those
taken in Wyoming. In the Colorado material, the fulvous color of the
limbal area of the secondaries is less pronounced and the general ap-
pearance of the underside is of a cooler tone of brown, that is, gray-
brown rather than a warm brown. These variations are not considered
as significant.

Acknowledgments

The authors would like to thank Dr. John Reeder, Curator, Rocky
Mountain Herbarium, University of Wyoming for making the determina-
tion of the host plant. Dennis R. Groothuis supplied the Converse Co.,
Wyoming collection records for schryveri. Additional Wyoming records
were taken from De Foliart (1956). M. Fisher and J. D. Eff provided
records and field assistance for the Colorado study. Thanks are also due

266 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

nd

John Emmel for making comments on this paper. It should be noted that
he has successfully reared schryveri larvae on Sedum spathulifolium
Hook., from eggs sent by Stanford and Eff. The botanical nomenclature
used herein is based on Porter (1964, 1968) and Craighead (1963).

Literature Cited

Brown, F. M. 1957. Colorado butterflies. Denver, Colo., 368 pp.

CRAIGHEAD, J. J., F. C. CraicHeAp, Jr., AND R. J. Davis. 1963. A field guide to
Rocky Mountain wildflowers. Boston, 277 pp.

Cross, F. C. 1937. Colorado butterflies. Proc. Colo. Mus. Nat. Hist., Vol. 16.

Der Fouiart, G. R. 1956. An annotated list of southeastern Wyoming Rhopalocera.
J. Lepid. Soc. 10: 91-101.

Porter, C. L. 1964. Wyoming trees. Circular 164R, Agricultural Extension Service,
University of Wyoming.

1968. Spring flora of southeastern Wyoming. Bulletin 491, Agricultural

Experiment Station, University of Wyoming.

LEREMA ANCILLARIS (HESPERIIDAE) NEW TO TEXAS
AND THE UNITED STATES

In an earlier paper (1970, Lepid. Soc. 24: 59), the author briefly described the
effects of hurricane “Bewlah” in 1967 on the Lepidoptera populations of extreme south
Texas, and the discovery of three Lycaenidae new to the U.S. A new skipper record
is now given, believed also to be associated with that hurricane:

Lerema ancillaris liris Evans, 1955. A catalogue of the American Hesperiidae indi-
cating the classification and nomenclature adopted in the British Museum (Natural
History). Part IV. Hesperiinae and Megathyminae. London. British Museum. 499
pp., pls. 54-88. (Describes new sub-species of Hesperiidae. Type locality, Atoyac &
Veracruz, Mexico. Type in B.M.).

One 6, good condition, Santa Ana National Wildlife Refuge, near Alamo, Hidalgo
Co., Texas, 28 September 1968 (leg. R.O.K.; det. H. A. Freeman).

Evans described liris from 21 specimens collected in Mexico, Guatemala, Roatan
Island, Honduras, Nicaragua, and Costa Rica. H. A. Freeman (personal communica-
tion) advises that this species is common over much of Mexico, and is one of the most
common species at Ciudad de Valles, Veracruz, and Oaxaca. Ciudad de Valles is
about 300 miles south of Santa Ana National Wildlife Refuge, site of the first U.S.
record.

Joseph F. Doyle, III records collecting a @ Synapte salenus Mabille, 27 August
1968 at the Santa Ana National Wildlife Refuge. No doubt the appearance of this
insect north of the Rio Grande was also associated with hurricane “Beulah.”

As with the Mexican Lycaenidae which were apparently displaced by “Beulah,” it
will be interesting to learn whether or not these Hesperiidae become established in
extreme south Texas. There are no other known records from the United States for
any of these species; however, other like specimens must have been present at the
time (1968). Should any of these become resident north of the Rio Grande, the sig-
nificance of great weather disturbances on insect dispersal will have been established.

Roy O. Kennay, 135 Vaughn Place, San Antonio, Texas.

VoLUME 24, NUMBER 4 267

A NEW SPHINX MOTH FROM THE WEST INDIES (SPHINGIDAE )
Mrs. C. Reep Cary!

Several years ago the late Mrs. C. Reed Cary prepared a manuscript in which the
following, interesting new sphinx moth was described. The publication of her paper
was withheld pending the completion of more extensive works, dealing with the
Macrolepidoptera of the West Indies, to which it was hoped this description could be
appended. Because the appearance of the larger papers will be delayed I take this
opportunity to present the following for the late Mrs. Cary.

Some editing of Mrs. Cary’s paper has been undertaken and drawings have been
prepared by Mr. Andre del Campo Pizzini, staff artist of the Smithsonian Institution.

J. F. G. Clarke
U.S. National Museum

In April, 1956, J. F. Gates Clarke, while a member of the Smithsonian—
Bredin Caribbean Expedition, collected five specimens of the genus
Cautethia at light on the small, isolated, and sparsely settled island of
Barbuda, in the Lesser Antilles. In 1958, while on another Smithsonian—
Bredin expedition he collected eight more of this moth on Antigua. The
13 specimens collected by Clarke differ so sharply from known forms that
I describe them as follows:

Cautethia noctuiformis bredini Cary, new subspecies
(Figures 2, 44a)

Alar expanse 28-34 mm.

Labial palpus white basally, suffused and irrorate grayish toward apex, the second
segment with large grayish fuscous blotch on outer side. Antenna gray; bases of
scales somewhat darker and tips of scales narrowly edged with sordid white. Head
gray, suffused fuscous; tips of scales white. Thorax gray, collar narrowly edged with
grayish fuscous; tegula with broad grayish-fuscous, longitudinal stripe. Forewing
ground color gray, antemedial line strongly outwardly curved, double, blackish fuscous;
postmedial line double, grayish fuscous; inner element outwardly curved to vein 5
then nearly straight to outer dorsum; outer element very strongly dentate to vein 2
then broadened to tornus; terminal line very narrowly dark gray; on apical third of
costa a dark gray triangle, its base on costa, extends one-third distance across wing;
cilia mixed gray and sordid white. Hind wing basal half mikado orange, terminal half
grayish fuscous; cilia mixed grayish fuscous and sordid white. Legs sordid white
speckled and suffused with grayish fuscous; tarsi marked with grayish fuscous. Abdo-
men gray dorsally, buff ventrally; fifth segment with grayish-fuscous blotch laterally.

Male genitalia (slides Cary No. 76; JFGC. No. 12080). Harpe broadest basally,
gently tapered to a narrowly rounded cucullus; sacculus moderately sclerotized, ter-
minating in a cupped process set with small teeth. Gnathos stout, enlarged distally and
divided postero-distally into two blunt, short, heavily sclerotized protuberances.
Uncus very stout terminating in a heavily sclerotized blunt point. Vinculum very

1 Deceased

268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figure 1. Cautethia noctuiformis noctuiformis (Walker )
Figure 2. Cautethia noctuiformis bredini, new subspecies

narrow; saccus produced as a linguiform process. Tegumen broad basally, narrowed
posteriorly. Anellus a broad band, narrowed laterally. Aedeagus longer than harpe,
slightly curved, dorso-apically produced as sharp point; vesica armed with two small
clusters of slender cornuti.

Holotype: U.S. National Museum No. 70895.

Type locality: British West Indies, Antigua, English Harbor.

Distribution: Antigua, Barbuda.

Described from the holotype (IV. 1958, J. F. G. Clarke), and 11 é paratypes from
Antigua with same data as holotype, and Barbuda, Oyster Pond (6. IV. 1956, J. F. G.
Clarke).

Unfortunately, no females of bredini were collected so only the male
genitalia can be illustrated for comparison.

In superficial appearance Cautethia noctuiformis noctuiformis (Walker )
and C. grotei grotei Edwards are similar species, but they are easily dif-
ferentiated by their genitalia (see Figures 3 and 4). C. n. bredini clearly
belongs with n. noctuiformis but can be distinguished from the typical
subspecies by its paler ground color and the more contrasting transverse
markings. The transverse markings of bredini are sharply defined (Fig-
ure 2) and the preapical dark gray triangle stands out in contrast to the
light gray of the forewing. On the other hand noctiuformis is strongly
suffused with very dark gray, the transverse markings are obscured and
the preapical triangle fuses with the dark coloring (Figure 1).

Our knowledge of the distribution of grotei and noctuiformis is too
fragmentary to permit generalizations, but for the time being we can state
that grotei (and its subspecies) is the dominant species in Cuba, Jamaica,

>

Migure 3. Cautethia grotei grotei Edwards: 3, Ventral view of male genitalia with
left harpe and aedeagus removed; 3a, aedeagus.

Migure 4. Cautethia noctuiformis bredini, new subspecies: 4, Ventral view of male

genitalia with left harpe and aedeagus removed; 4a, aedeagus.

VoLuME 24, NuMBER 4 269

270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Cayman Islands and Hispaniola, and noctuiformis is dominant on Puerto
Rico, Virgin Islands, and a few of the Lesser Antilles.

It gives me great pleasure to name this subspecies for Mr. and Mrs.
Bruce Bredin who sponsored the expedition on which the specimens were

collected.

A RECORD OF A NATURALLY OCCURRING LIMENITIS HYBRID
(NYMPHALIDAE )

A suspected hybrid between Limenitis archippus Cramer and Limenitis lorquini
Boisduval was collected two miles south of the Richland Y on Amon wasteway, Benton
County, Washington. On July 6, 1969, three further hybrids were caught at the junc-
tion of Amon wasteway and the Yakima River by J. C. Montgomery.

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Left: Limenitis archippus. Centre: Suspected hybrid between Limenitis archippus
and L. lorquini. Right: Limenitis lorquini.

The four specimens of the Limenitis hybrids vary in color and markings. Limenitis
archippus and lorquini both fly in this locality and they are the only species of the
genus Limenitis in the area.

Epwarp V. Gace, Route 1, Box 10, Richland, Wash.

VOLUME 24, NuMBER 4 PATA

LARVAE OF HAPLOA CLYMENE (ARCTIIDAE) HIBERNATING
ON CONCRETE BRIDGES

Larvae of the arctiid, Haploa clymene Brown, have been found on concrete bridges,
sites used for collecting winter stoneflies from northern Illinois to Florida. The larvae,
encountered on numerous occasions, were often found on the completely exposed
surfaces of the pillars, railings, etc., but more commonly in cracks, between pillars,
and such places. The numbers on a bridge varied from one or two up to 50 or so.
Larvae collected throughout the winter appear to be in the same stage with a length
of 15-20 mm. Caterpillars were found from November through March.

Subsequent observations revealed that the larvae were feeding on small willows
which grow along streams in central Alabama. Feeding was seen as late as November.
In early November on one stream, apparently hibernating larvae were found on large
rocks upstream from the bridge. It appears that the caterpillars also migrate onto
bridges to hibernate since this is similar to the rock habitat.

Fig. 1. Map of Alabama. Localities where larvae of Haploa clymene Brown were
taken on concrete bridges are indicated by dots.

Hibermating larvae taken from a bridge on November 7, 1968 in central Alabama
were reared on an artificial diet (Ann. Ent. Soc. America 62: 1216) in our laboratory.
After one week they became active, began to eat, and molted soon thereafter. They
continued to grow slowly, reaching a maximum size of about 35-40 mm. Pupation
occurred on December 28, 1968, followed by emergence on March 30, 1969.

It is highly probable that such hibernation occurs in other areas within the range of
Haploa clymene. Specific collection localities in Alabama are indicated by dots on the
accompanying map. This species appears to range throughout the state. Interested
persons should check their area for the “concrete bridge” caterpillar.

T. D. Ercan ann H. B. CunnincuaM, Dept. of Zoology-Entomology, Auburn Uni-
versity, Auburn, Alabama.

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JOURNAL OF THE LEPIDOPTERISTS SOCIETY

CANADIAN SPECIES OF LITHOCOLLETIS FEEDING ON
SALIX AND POPULUS (GRACILLARIIDAE)

T. N. FREEMAN

Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario

This paper defines the known Salix and Populus feeding species of
Lithocolletis in Canada and the United States of America, in order that
other workers may present their accumulated data on life history and
ecology. The study was based mainly on material collected by officers
of the Canada Forest Insect Survey and on that reared by me and my
associate, Mr. G. G. Lewis. All existing types of described North Ameri-
can species were examined. Diagnostic characters of the male genitalia
and maculation are presented. I have been unable to associate any species
with Lithocolletis salicifoliella Clemens (1861, Proc. Ent. Soc. Philadel-
phia 1: 86) which was described on the basis of the larval mine. The
mine is on the under surface along the edge, and usually near the base
of the leaf of Salix alba Linn. Until adult material is obtained from near
the type locality at Easton, Pennsylvania, the species cannot be recog-
nized.

Key to Species

1. Valvae symmetrical, terminating in a downward curving, pointed apex (Figs.

9, 12). Larva mining the leaves of Populus spp. = ee 2
Valvae asymmetrical, or if symmetrical, terminating in a knob of short spines.
(Figs. 14, 15). Larva mining the leaves of Salix spp, =e 6
2. Larva mining the leaves of Populus balsamifera L. _-.-- nipigon sp. n.
Larva mining the leaves of other Populus spp, —- 2 eee 3
3. Thorax of adult shining ochreous or golden, with a narrow, white, median,
longitudinal ‘streak ceo. see ee ee ee populiella Chamb.
Thorax not as above: .2.0022o 2 ee 4
4. Head and thorax entirely white. Forewing predominantly white ___ atomariella Zell.
Head, thorax, and forewing not as above 9 5
5. Forewing with four white streaks from the trailing margin, the fourth small
and near the wing apex. Wingspread 6.5-8.0 mm. __....._.__-___-___ ontario sp. n.
Forewing with five white streaks from the trailing margin, the fifth small and
near the wing apex. Wingspread 9.0-10.5 mm. _......-__-___ tremuloidiella Brn.
6. Forewing with three white streaks from the trailing margin _____ salicivorella Brn.
Forewing with four white streaks from the trailing margin, the fourth small
and near the wing apex _:... 2 ee i,
7. Forewing with four white streaks from the costal margin __ scudderella F. and B.
Forewing with five white streaks from the costal margin ___._______ kenora sp. n.

Lithocolletis atomariella Zeller

Zeller, 1875. Verh. Zool.-bot. Ges. Wien. 25: 350.

Maculation (Fig. 1). The following is a translation of Zeller’s original description:
rhorax and head white; tuft divided into two divergent tufts, also whitish, some

VoLUME 24, NuMBER 4 DHE

of the hairs sometimes with brownish tips. Antennae white, more or less distinctly
ringed. First segment of hind tarsus white, apical third brownish grey; the following
three segments brownish grey with pure white bases; apical segment whitish. Ab-
domen dark grey with whitish anal segment.”

“Width of forewing not entirely constant, but broader than in comparella and
narrower than in populifoliella, without shine, pure white, with a few, black, tiny
particles. The markings consisting of clay-yellow or dark ochre coloured spots which
have posterior black margins. There are five on the costal margin; the first two are
very oblique, sharply pointed; the first is elongated along its costal margin toward
the base; the third is less oblique, and the following two, which are greatly reduced
in size, are almost vertical; the last smallest one is extended in a curve to the hind
margin and encloses the black apical streak. Near the base of the hind margin is a
little spot which is often small or obscure, and is sometimes separated from the
margin. The first dorsal spot is very oblique, sharply triangular, extending to the
apex of the first costal spot, and forms with it a sharply angled broken line. The
second dorsal spot, which is widely separated from the first, is longer and variable
in size, and unites with the third and fourth costal spots, and since the third dorsal
spot is inclined inwardly and also unites with the second, a somewhat X-like figure
is formed with its apices extending to opposite margins. The black apical streak is
short and margined below and apically with clay-yellow, above it the white ground
colour appears as a costal spot (apically from the fifth yellow costal spot). At the
base of the whitish apical fringe is a blackish curved line which reaches almost to
the third dorsal spot.”

“Hind wing grey with paler fringe. Underside of forewing dark grey with four
indistinct costal spots which increase in size apically.”

“Habitat: Cambridge, Massachusetts (Hagen) where this species is not often seen.”

Male genitalia of type (Fig. 9). Symmetrical. Valva long, narrow, arcuate dor-
sally; apical portion obtusely angled, recurved and pointed; inner surface with nu-
merous, short, thick spines. Uncus conical. Transtilla arched. Aedeagus tubular,
weakly sclerotized. Flap of eighth sternite spatulate; apex invaginated.

Remarks. I have not seen any specimen that will match Zeller’s type.
No food plant was given in the original description, but because other
species with this type of genitalia feed on Populus, I assume that is the
food plant. The type is in the U.S. National Museum, Washington, D.C.
The species is allied to L. tremuloidiella as judged from the similarity of
the male genitalia.

Lithocolletis ontario Freeman, new species

Maculation (Fig. 2). Antenna with alternating fuscous and white bands. Face
white. Tuft on vertex fuscous beneath, white above. Thorax white, mottled with
black. Forewing with white and golden ochreous markings, the former speckled with
fuscous and the latter narrowly margined with black apically. Four dorsal and five
costal white streaks. Base of forewing white extending narrowly over the fold to
the apex of the first rectangular dorsal, white streak; the first costal streak extends
obliquely outward to the fold where it is narrowly connected to the second dorsal
streak, situated near the middle of the wing, and narrowly extended apically to join,
or almost join, the second and third costal streaks; the third dorsal streak triangular,
its apex extending about half way across the wing and situated opposite the space
between the third and fourth costal streaks; the fourth costal and fourth dorsal streak
usually confluent and the golden ochreous markings between them and the first dorsal
and second costal streaks suggesting the letter K; the fifth costal streak short and in-
clined inwardly; a small, black spot, bordered below with a few white scales near

O74 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

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VoLUME 24, NuMBER 4 RI:

apex; apex with a curved row of dark fuscous scales; fringe whitish. Hind wing light
fuscous; fringe whitish. Wingspread: 7—8 mm.

Male genitalia (Fig. 10). Symmetrical. Valva narrow, tapering; apical portion
decurved in variable degree; sparsely covered with fine spines. Vinculum V-shaped.
Transtilla a semi-circular band. Uncus long, conical. Anellus thick, tubular. Aedeagus
short; apex hooked. Flap of eighth sternite spatulate; apex usually invaginated.

Flight period. August and early September.

Distribution. Known only from many localities in southern and northwestern On-
tario, but undoubtedly more widely distributed.

Food plants. Populus tremuloides Michx. and P. grandidentata Michx.

Type material. Type male, Simcoe, Ontario, 5 Sept. 1957 (Freeman & Lewis),
rearing lot No. 57-174 from Populus tremuloides Michx. No. 10,890 in the Canadian
National Collection, Ottawa, Ont. Paratypes: Nine males, six females, Simcoe, Ont.,
5-11 Sept. 1957 (Freeman & Lewis), rearing lot number and host same as those
of type. One male, two females, Simcoe, Ont., 9-13 May 1957 (Freeman & Lewis),
rearing lot No. 57-172 from Populus grandidentata Michx. One male, Simcoe, Ont.,
13 Sept. 1965 (Freeman & Lewis), rearing lot No. 65-83 from Populus tremuloides.

Mine. A blotch on the underside of the leaf.

Remarks. The maculation of this species resembles that of Lithocolletis
atomariella Zell. but it has many more fuscous scales in the white areas
of the forewing, and much finer spines on the valva of the male genitalia.
The male genitalia of ontario resemble those of L. tremuloidiella Brn.
(Fig. 11) but the maculation differs. There are four dorsal white streaks
on the forewing of ontario and five on that of tremuloidiella. Ontario is
a Huron Indian word meaning sparkling water.

Lithocolletis tremuloidiella Braun

Braun, 1908. Ent. News 19:. 102.

Maculation (Figs. 3, 4). The following is Dr. Braun’s original description:

“Antennae dark gray, the joints becoming lighter toward their bases. Palpi grayish
white. Face grayish white. Tuft gray, mixed with white. Forewings pale reddish
brown near the base, becoming more ocherous beyond the middle. There is a short
median basal white streak, and a dorso-basal white streak, both thickly dusted with
blackish scales, and uniting with the first dorsal streak. There are five costal and
five dorsal white streaks, all dark margined internally and more or less dusted with
blackish scales, the last two pair, however, not dusted. These streaks are situated as
follows: a large oblique white costal streak at the basal fifth, attaining the middle
of the wing and prolonged backward; nearer the base an oblique dorsal streak, almost
meeting the first costal before the middle; the second costal streak also oblique; the
' second dorsal streak nearer the base than the second costal, large, oblique, strongly
constricted just below the fold, and uniting with the apices of the first and second
costal streaks; third costal and third dorsal streaks nearly opposite, less oblique, their
apices separated by a narrow prolongation of the blackish internal margin. The

&

Figs. 1-8. Adults of Lithocolletis spp. 1, Male type, L. atomariella Zell., Cam-
bridge, Mass.; 2, male type, L. ontario sp. n., Simcoe, Ont.; 3, 4, female type and
male paratype, L. tremuloidiella Brn., Kaslo, B.C.; 5, male type, L. nipigon sp. n.,
Fort William, Ont.; 6, male type, L. salicivorella Brn., Essex Park, N.J.; 7, male, L.
populiella Chamb., Cincinnati, Ohio; 8, male type, L. kenora sp. n., Ramsay, Ont.

976 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

fourth costal and dorsal streaks, of which the costal is at the apical fourth, the dorsal
at the tornus, form an inwardly convex white fascia. Fifth costal and dorsal streaks
also form a curved white fascia. An irregular blackish apical spot. Marginal line in
the cilia blackish with a bluish luster. Cilia gray. Alar expanse 9-10.5 mm. Hind
wings gray. Cilia gray with a reddish tinge. Abdomen dark gray above, grayish
white beneath. Anal tuft grayish ocherous. Legs dark gray, tips of the tarsi lighter.”

“Described from specimens bred by Mr. W. D. Kearfott from tentiform mines on
the underside of leaves of Populus tremuloides Michx., received from Mr. J. W.
Cockle, Kaslo, B.C., Aug. 26, 1907. The imagos appeared during the same month.
The mine is large (for an underside Lithocolletis mine), oval, and with a fine fold
through its long axis... .”

Male genitalia of paratype (Fig. 11). Symmetrical. Valva long, narrow, arcuate
dorsally; apical portion recurved, pointed, almost right angled; inner surface with
several, short, thin spines. Uncus conical. Transtilla arched. Aedeagus tubular.
Flap of eighth sternite spatulate.

Remarks. This species is known to me only from Dr. Braun’s type
and paratype in the U.S. National Museum, Washington, D.C. The
species is allied to L. ontario as judged from the similarity of the male
genitalia.

Lithocolletis nipigon Freeman, new species

Maculation (Fig. 5). Antenna with alternating grey and white bands. Face white.
Tuft on vertex fuscous beneath, white above. Thorax mottled with black and white.
Forewing with five costal and four dorsal white streaks dotted with fuscous, and
with brownish-yellow markings narrowly margined with black apically as follows:
one at basal quarter extending from costa obliquely outward about half way across
wing; one before middle of costa extending obliquely outward about half way across
wing, then angling backward as a short, oblique spur; a costal spot just beyond
middle; a somewhat outwardly oblique fascia before apical region; a patch heavily
margined with black apically near trailing margin, between and posterior to the first
two costal markings; a larger patch from just beyond middle of trailing margin, ex-
tending obliquely outwardly to middle of wing, and often uniting with subapical
fascia. Apical region pale grey with black-tipped scales sometimes united to form
a black spot. Fringe pale fawn with a narrow, black, arcuate streak. Hind wing
and fringe pale grey. Wingspread 8-9 mm.

Male genitalia (Fig. 12). Symmetrical. Valva long, narrow, with apical portion
pointed, curved; inner surface with thin, short spines. Vinculum conical. Transtilla
a semicircular band. Uncus very long, conical. Aedeagus long, tubular, weakly
chitinized. Flap of eighth sternite spatulate, usually tapering; apex invaginated.

Type material. Holotype: male, Fort William, Ontario, 20 Aug. 1962. Forest In-
sect Survey No. S62-5112-01. No. 10,888 in the Canadian National Collection, Ot-
tawa, Ont. Paratypes: six males, six females, Kivikoski, Ontario, 20-27 Aug. 1962;
five males, three females, Fort William, Ontario, 20 Aug. 1962; one male, one female,
Sioux Lookout, Ontario, 2 Aug. 1962 and 23 July 1962 respectively; one male, Ono

>

Figs. 9-16. Male genitalia of Lithocolletis spp. 9, Type, L. atomariella Zell.,
Cambridge, Mass.; 10, paratype, L. ontario sp. n., Simcoe, Ont.; 11, paratype, L.
tremuloidiella Brn., Kaslo, B.C.; 12, paratype, L. nipigon sp. n., Kivikoski, Ont.;
13, type, L. scudderella F. & B., probably Boston, Mass.; 14, paratype L. kenora sp. n.,
Shebandowna, Ont.; 15, type, L. salicivorella Brn. Essex Park, N.J.; 16, L. populiella
Chamb., Cincinnati, Ohio.

VoLuME 24, NuMBER 4 OCT

278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Park, Ontario, 22 Aug. 1962. All type material reared by officers of the Forest Insect
Survey, Canada Department of Fisheries and Forestry from Populus balsamifera L.
Distribution. Known at present only from western Ontario but undoubtedly more
widely distributed.
Mine. A blotch on the underside of the leaf.

Remarks. This species has not been found on species of Populus other
than the one above. There is only one generation each year. As judged
by the male genitalia, this species belongs in the L. atomariella Zell.
group, but nipigon is easily recognized by the gentle downcurved valva
apex. The place name Nipigon comes from a Cree Indian word mean-
ing the lake you cannot see the end of.

Lithocolletis scudderella Frey and Boll

Frey and Boll, 1873. Stett. ent. Zeit. 34: 212.

Maculation. The following is a translation of Frey and Boll’s original description:

“Of medium size; however, several specimens are considerably smaller; rather
broad-winged. The ground color of the head, thorax and forewings is a peculiar
light yellowish brown. This color is similar to that of a flown specimen of L. con-
nexella Z. The forewings are finely scaled. Head and palpi brownish white. Antennae
brownish white, annulate with darker. The legs are brownish white, the tibiae of
the first pair are striped with black. Thorax concolorous with the wings, with a
median whitish line. Patagia whitish. Abdomen gray, lighter at the tip.”

“The markings of the forewings are somewhat indistinct, but are characteristic
because of the great accumulation of black scales in the region of the fold. They are
as follows: there is a rather straight basal streak ending at one-third the wing length,
where it is slightly dilated. It is margined with blackish around the apex. The first
pair of streaks are placed very obliquely. Of these, the costal is the smaller and is
triangular. The dorsal is of very peculiar form, in that it is strongly constricted on
the fold. Both are black margined internally. Usually the extent of the black scales
on the inner edge of the dorsal streak is so greatly increased as to form a considerable
sized black spot, which extends to the apex of the basal streak. (Less frequently
these black scales are almost lacking). The next pair of streaks are at the middle
of the wing length, and are also inwardly black margined. (These black scales are
also increased so as to form a spot). The costal streak is narrow, almost perpendicular;
the dorsal forms a broader triangle, placed just before the tornus. Toward the apex
are two very narrow, slightly curved streaks. At the apex is a black dot or a streak
bordered with white scales toward the base. Cilia light, darker toward the tornus,
with a blackish marginal line around the apex.”

“Hind wings dark gray. Cilia somewhat lighter.”

“The last three light costal streaks are visible on the dark brownish underside of
the forewings.”

“The larvae are abundant in October and November on several species of Salix.
The mine is on the underside between the veins. It is elongate-oval and somewhat
wrinkled. The imagoes emerged in April and May.”

Male genitalia of type (Fig. 13). Asymmetrical. Right valva elongate, slightly
tapering; apex notched; a recurved, subapical, ventral spine. Left valva broad, sub-
ovate; apex with stout, recurved, ventral spine. Uncus broad, subovate. Flap of
eighth sternite broad, spatulate (damaged).

temarks. This species is known to me only from the original descrip-
tion, and the male genitalia of the type, that apparently was collected
near Boston, Mass. The type is in the British Museum (Natural History),

VoLUME 24, NuMBER 4 279

London, England. As judged from the male genitalia, the species is
allied to L. salicivorella Brn.

Lithocolletis salicivorella Braun

Braun, 1908. Ent. News 19: 101.

Maculation (Fig. 6). Dr. Braun’s type in the United States National Museum is
somewhat faded. I therefore quote her original description:

“Antennae pale grayish ocherous, faintly annulate with darker, somewhat darker
at the tip. Palpi shining white. Face white. Tuft pale gray with an ocherous tinge.
Thorax and basal third of the forewings ocherous gray. Wings becoming more ocher-
ous towards the apex. A white band across the anterior margin of the thorax extends
across the patagia and is continuous with a median basal white streak on the fore-
wings. There is a short dorso-basal white streak, somewhat dilated posteriorly. The
median basal streak is curved downwards, and extends for two-fifths of the wing
length, where it is confluent with the first dorsal streak, its upper edge uniting with
the apex of the first dorsal streak. A few dark brown scales extend around the apex
of the angle thus formed. There are four costal and three dorsal white streaks. The
first dorsal streak at the basal fourth is very large, oblique and curved. Internally,
it is dark margined just before it unites with the basal streak, the dark margin being
continued around the angle and for a short distance along the lower side of the
basal streak. The extreme edge of the costa is dark brown for about one-third of
the wing length, where the dark line is deflexed, and continues as the dark margin
of the first costal streak which is narrow, very oblique, its apex extending to a point
just beyond the apex of the first dorsal. The second costal streak is large, nearly
perpendicular, its apex opposite to that of the second dorsal streak, which is also
very large, and placed slightly nearer the base than the corresponding costal streak,
and is somewhat oblique. The next pair of streaks, of which the dorsal is placed
just above the tornus, are nearly opposite to each other, slightly oblique toward the
base and eurved; their apices nearly meet. These two pairs of streaks are margined
internally with brown scales. Fourth costal streak oblique, curved and unmargined.
A narrow line of black scales extends from below the apex of the last costal streak
to near the apex of the wing, and is margined above by a line of white scales. Mar-
ginal line in the cilia dark brown, extending around the apex from the fourth costal
streak to the third dorsal. Cilia grayish ocherous. Alar expanse 7 mm. Hindwings
grayish. Cilia gray, with a fulvous tinge. Abdomen dark gray above, whitish beneath.
Anal tuft grayish ocherous. Legs whitish, banded and striped with gray.”

“The unique type, a male of this species, was bred by Mr. W. D. Kearfott from a
much wrinkled mine on the underside of a willow leaf, collected in Essex County
Park, N.J., July 6, 1902. The imago appeared July 19.”

“This species is nearest to L. scudderella Frey & Boll, which it closely resembles
in the costal and apical markings.”

Male genitalia of type (Fig. 15). Asymmetrical. Right valva long, narrow, with
a sub-apical, long ventral spine. Left valva sub-ovate, with a long, curved, stout,
ventral spine. Uncus long, conical. Saccus U-shaped. Aedeagus tubular; apex barbed.
Flap of eighth sternite broad, gradually tapering toward apex.

Distribution. Known to me only from the type locality, Essex Park, New Jersey.

Remarks. On the basis of the male genitalia, this species is allied, as
Dr. Braun states, to L. scudderella Frey & Boll.

Lithocolletis populiella Chambers

Chambers, 1878. Bull. U.S. Geological Survey 4: 101.

Maculation (Fig. 7). Antenna, head and legs white. Thorax pale golden, with
median, and two lateral, longitudinal, white streaks. Abdomen pale golden above.

280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Forewing pale golden with basal costal, median, and posterior white streaks, all ex-
tending to basal one-third; four costal and three dorsal white streaks dark margined
basally; first costal and dorsal ones extending obliquely outward; second and third
costal ones almost perpendicular, the second much smaller; second dorsal extending
obliquely outward to middle of wing, triangular; third dorsal almost perpendicular
to middle of wing; fourth costal near apex, large, rectangular; fringe white, with
brownish basal line. Hind wing fuscous; fringe white. Wingspread 7.5-8.5 mm.
Legs white.

Male genitalia (Fig. 16). Asymmetrical. Valva broad; apical portion with thick
spines; left valva with curved apical spine. Uncus long, conical. Saccus U-shaped.
Aedeagus tubular; apex barbed. Flap of eighth sternite subtriangular.

Distribution. Known to me only from Kentucky and Ohio.

Mine. A small tentiform underside blotch on the leaves of Populus alba L.

Remarks. On the basis of the male genitalia, this species belongs to
the L. atomariella Zell. group. Chamber’s type in the Museum of Com-
parative Zoology, Cambridge, Mass., consists of only the forewings. My
description was made from specimens from Cincinnati, Ohio. According
to Braun (1908, Revision of the North American species of the genus
Lithocolletis, p. 304) “the pupa is not enclosed in a cocoon, but its anal
end is attached to a small button of silk toward one end of the roof of
the mine.”

Chambers described this species from specimens from Kentucky and
Ohio. Apparently, the only remaining specimen of his material, is one
in the Museum of Comparative Zoology. This consists of both forewings
glued to a pin and bears the following three labels: “type 1510”; “122”;
“Kentucky.” This specimen, which I have examined, I designate as lecto-

type.

Lithocolletis kenora Freeman, new species

Maculation (Fig. 8). Antenna with alternating brown and white bands. Face glossy
white. Tuft on vertex white, mixed with dark brown. Thorax white, sometimes with
a few, dark-brown speckles. Forewing white with glossy fawn markings, edged
apically with black, and with five costal and four dorsal white streaks sometimes
containing a few, scattered, blackish scales. The glossy fawn markings arranged as
follows: a small, often elongate one in anal region; an outwardly oblique fascia ex-
tending from basal one-quarter of costa to middle of wing, then abruptly angled to
trailing margin, sometimes with a narrow band of white separating costal and dorsal
portions; another oblique fascia extending from costal half to above fold, where a
somewhat anteriorly curved spur extends to a K-shaped fascia located near apical
three-quarters; at apex a small patch, almost enclosing a small black dot, and with
a curved spur extending to costa; apical patch markedly edged with a curved black
aS Fringe whitish grey. Hind wing fuscous with lighter fringe. Wingspread

—9 mm.

Male genitalia (Fig. 14). Symmetrical. Valva broad; dorsal region with a large
patch of spines; an area of about ten scattered spines near middle of ventral margin;
a spinose knob and a blunt hook on ventral margin, near apex; apical margin some-
times with a blunt tooth. Uncus long, conical. Transtilla rectangular. Saccus broadly
V-shaped. Aedeagus long, tubular, with a long, chitinized supporting rod, and barbed
apex. Flap of eighth sternite spatulate.

Type. Male, Ramsay, Ont., 3 Aug. 1962, No. 10,889 in the Canadian National

VoLuME 24, NuMBER 4 281

Collection, Ottawa. Paratypes: two males, three females, Red Lake, Ont., 15 Aug.
1962. One male, Shebandowan, Ont., 9 Aug. 1963. One male, one female, English
River, Ont., 23-25 Aug. 1962. One male, Ramsay, Ont., 3 Aug. 1962. One female,
Ignace, Ont., 29 Aug. 1962. One female, Finland, Ont., 17 Aug. 1962. All type
material seared from Salix spp. by officers of the Forest irasect Survey, Canada De-
partment of Fisheries and Forestry.

Distribution. Known only from western Ontario, but undoubtedly more widely
distributed.

Mine. A blotch on the underside of the leaf.

Remarks. Clemens (1861, Proc. ent. Soc. Philadelphia 1: 81) de-
scribed Lithocolletis salicifoliella from a mine found on Salix alba L. at
Easton, Pennsylvania. He reared no adults but stated: “The mine is on
the undersurface, usually near the base of the leaf and along the edge.”
Until specimens are reared from Salix at Easton, it is impossible to apply
the name salicifoliella to any known species. As judged by the male
genitalia, kenora is closely related to the Rubus feeding Japanese species
L. pulchra Kumata, which possesses a median, basal, white streak. This
is absent in kenora.

The place Kenora used to be called Rat Portage. It was changed to
Kenora, a word derived from the first two letters of the following: Kee-
watin, Norman River and Rat Portage.

Acknowledgments

I am indebted to the following persons who have assisted me by lend-
ing types, or providing information on them: Dr. A. F. Braun, Cincinnati,
Ohio; Drs. D. R. Davis and R. W. Hodges, United States National
Museum, Washington, D.C.; Messrs. P. E. S. Whalley and A. Watson,
British Museum (Natural History), London, England; Dr. P. J. Darling-
ton, Jr., Museum of Comparative Zoology, Cambridge, Mass.; Mr. O. H.
Lindquist and others, Forest Insect Survey, Canada Department of
Fisheries and Forestry, Sault Ste. Marie, Ontario. Finally, I appreciate
the assistance of Mrs. A. Mutuura, Ottawa, Ontario, who prepared most
of the coloured illustrations accompanying this paper.

282, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

THE LIFE HISTORY OF SCHINIA FLORIDA (NOCTUIDAE)

D. F. HARDWICK

Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario

Schinia florida (Guenée 1852) feeds in the larval stage on the Evening
Primrose, Oenothera biennis L. Previous notes on its life history have been
published by Fitch (1868), Saunders (1869), Kellicott (1879), and
Beutenmuller (1901), and the ultimate-stadium larva has been described
by Crumb (1956).

The species is widespread in temperate North America, being distrib-
uted in southern Canada and the northern United States from the Atlantic
coast to Alberta and eastern Washington. In the East, it occurs southward
to North Carolina and in the West, to Colorado and Utah. It evidently
does not occur in the extreme southwestern United States; the latter area
is inhabited by the closely related Schinia felicitata (Smith), which feeds
in the larval stage on the white-flowered Oenothera deltoides Torr. (for
life history data, see Hardwick 1967).

In different parts of its range, Schinia florida may be found in flight
between the end of May and the first of September. Because eggs are laid
on the buds, the seasonal period of adult activity is co-ordinated with the
blossoming period of the food plant. In the Ottawa area, the moth is
usually common throughout the months of July and August. The rather
protracted flight period led Forbes (1954) to suggest that the species may
be at least partially double-brooded, but there is no clear evidence of this.
None of the specimens reared in conjunction with the present study
emerged in the same year that they pupated.

Behaviour

Schinia florida is an exclusively nocturnal species and becomes active
only at dusk. During the hours of daylight, the moth rests on the flowering
stalk of its food plant. Most frequently it enters the cup-shaped corolla at
night and the petals actually close over the moth with the coming of day-
light (Fig. 1). Usually only the yellow apical quarter of each forewing is
left protruding beyond the petals, and these in their yellow colouring
closely resemble the petals themselves. This behaviour pattern is not an
absolute one, however; not infrequently the little moth merely nestles
among the blossoms, and the pink of the basal three-quarters of the wings
so closely simulate the colour of the dead but still clinging petals of
Oenothera that even in this relatively exposed position, the moth is still
difficult to detect.

VoLUME 24, NuMBER 4 283

Figs. 1-6. Schinia florida (Guenée). 1, Adult resting in blossom of Oenothera
biennis L.; 2, adult, Alberton, P.E.I.; 3, dorsal aspect of ultimate-stadium larvae; 4,

larva feeding on buds of Oenothera biennis; 5, left lateral aspect of ultimate-stadium
larvae; 6, pupae.

The eggs are laid on the buds at the apex of the flowering shoot of the
food plant; no effort is made to insert the eggs between the buds nor to
conceal them beneath the petals. Five individually confined, wild-caught
females deposited a mean of 114 eggs, and the maximum deposited by a
single female was 198.

284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The newly hatched larva bores directly into one of the small buds and
feeds on the sexual organs of the plant. As the larva increases in size,
successive buds are attacked in a similar fashion. Larger larvae also
attack the younger and more tender seed capsules by boring a hole
through the wall to gain access to the developing seeds. During the later
stadia, the larva does not usually secrete itself within a bud or seed cap-
sule but feeds from an exposed position on the stem (Fig. 4); it evidently
relies on its resemblance to the buds or seed capsules to protect it from
predators. The skin of the larva is densely set with elongate spinules
which give it a pubescent appearance similar to that of its food plant. In
the last stadium, larvae exhibit two colour phases, one green and one dull
red. The dull-red colour phase is evidently dependent for protection on its
resemblance to dying petals or to those areas of plant tissue with a red
suffusion that are characteristic of Oenothera biennis.

The fully grown larva tunnels into the ground to form its pupal cell, and
it is in the pupal stage that the insect overwinters.

Description of Stages

The descriptions of immature stages presented here were based on the
progeny of five females taken in the Ottawa area. Larvae were reared
individaully at room temperature on the buds and seed capsules of
Oenothera biennis. Rearing methods employed were those outlined by
Hardwick (1958). The estimate of variability following the mean for
various values is the standard deviation.

Adult (Fig. 2). Head and prothorax pink. Pterothorax pale yellow dorsally.
Abdomen pallid yellow or creamy-white dorsally. Undersides of thorax and abdomen
pink, or pale yellow suffused with pink. Forewing pink marked with light yellow. A
quadrate yellow patch in posterior area of basal space. Remainder of basal space,
median space and subterminal space light to dark pink. Median space often a paler
pink than basal and subterminal spaces. A pale transverse posterior line, and darker-
pink orbicular and reniform spots, often evident. Subterminal line irregular, strongly
outlined by colour change between subterminal and terminal spaces. Terminal space
light yellow. Fringe concolorous with, or somewhat darker than, terminal space.
Hind wing uniform creamy white. Fringe concolorous. Underside of both wings pale
yellow, suffused with pink along costal margins. Fringes pale yellow.

Expanse: 31.2 + 1.6 mm (60 specimens ).

Egg. Prominently ribbed on micropylar and lateral surfaces; rather deep yellow on
day of deposition. A broad, subequatorial, orange girdling band becoming evident on
micropylar half of egg on the day after deposition. Girdling band darkening to a dark
red, and remainder of egg darkening to a yellow-orange by day of hatching. Blackish
head capsule of larva becoming visible through chorion of micropylar surface a few
hours before hatching.

Dimensions of egg: length, 0.651 + 0.048 mm; diameter, 0.713 + 0.039 mm (234
C2ZRS).

Incubation period: 4.3 + 0.6 days (313 eggs).

Virst-Stadium Larva. Head dark blackish-brown or black. Prothoracic and suranal

VoLUME 24, NuMBER 4 285

Figs. 7, 8. Schinia florida (Guenée), apical abdominal segments of pupa. 7, Ven-
tral; 8, right lateral.

shields dark brown, somewhat paler than head. Trunk cream, yellow, or yellowish-
green; a pair of subdorsal reddish-orange lines and occasionally a middorsal orange
line becoming evident in older larvae. Rims of spiracles and thoracic legs dark brown.

Head width: 0.392 + 0.009 mm (25 larvae).

Duration of stadium: 3.2 + 0.8 days (68 larvae).

Second-Stadium Larva. Head light to medium orange-brown; in some specimens
dorsal half of head mottled with darker brown. Prothoracic shield varying from
yellow-fawn to orange-fawn, variably mottled with brown; shield in some specimens
emarginated laterally and posteriorly with darker brown. Suranal shield straw yellow,
often with a greenish suffusion. Middorsal band greenish-grey or yellowish-grey.
Subdorsal area pale green or dull yellow, margined outwardly with a dark-pink line.
Lateral and ventral areas of trunk varying from pale green to dull greyish-yellow. Setal
bases light brown. Spiracles with medium brown rims. Thoracic legs varying from
light to medium brown.

Head width: 0.614 + 0.042 mm (25 larvae).

Duration of stadium: 2.1 + 1.1 days (68 larvae).

Third-Stadium Larva. Head pale green, pale fawn or pale orange-yellow; dorsal
half of head with faintly darker mottling. Prothoracic shield concolorous with head,
often flushed laterally with dark pink. Suranal shield pale yellow or pale green. Mid-
dorsal band dull green or dull grey. Subdorsal area paler than middorsal band, pale
green or straw-yellow; margined laterally by an indistinct inner greyish-white line and
an outer dark-pink line. Supraspiracular area essentially concolorous with subdorsal
area. Spiracular band a paler yellow or green than supraspiracular area. Suprapodal
area similar in colour to supraspiracular area but with a strong greyish tone. Mid-
ventral area paler than suprapodal area. Spiracles with light brown rims. Setal bases
essentially concolorous with trunk. Thoracic legs pale green or pale straw-yellow.

Head width: 0.985 + 0.076 mm (25 larvae).

Duration of stadium: 2.3 + 0.9 days (68 larvae).

Fourth-Stadium Larva. Head ochre or greenish-ochre, without any darker mottling.
Prothoracic shield greenish-ochre, often flushed with dark pink laterally. Suranal
shield light green. Trunk green, variably suffused with pink. Middorsal band narrow,
dark green. Subdorsal area green, paler than middorsal band; margined laterally by an
inner greyish-white line and an outer dark-pink line. Supraspiracular area concolorous
with subdorsal area. Spiracular band pallid green or greenish-yellow. Suprapodal area

286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

concolorous with subdorsal area. Midventral area light greyish-green. Spiracles with
light-brown rims. Setal bases concolorous with trunk. Thoracic legs light green.

Head width: 1.65 + 0.09 mm (25 larvae).

Duration of stadium: 3.6 + 0.9 days (68 larvae).

Fifth-Stadium Larva (Figs. 3, 5). Occurring in a green and a dull-red colour
phases, the former by far the commoner.

Green colour phase: Head pale yellowish-green. Prothoracic shield yellowish-green
variably suffused with purplish-pink, often heavily so. Suranal shield light yellowish-
green, essentially immaculate. Middorsal band narrow, dark green. Subdorsal area a
lighter green than middorsal band, often with yellowish segmental patches; subdorsal
area margined laterally by a greyish-white line and a purplish-pink band; pink band
evanescing in some cases. Supraspiracular area concolorous with, or somewhat darker
than, subdorsal area. Spiracular band pallid greyish-yellow, often poorly distinguished
from suprapodal area. Suprapodal area essentially concolorous with supraspiracular
area. Midventral area pale green with a powdery-grey suffusion. Spiracles with light-
brown rims. Setal bases concolorous with trunk. Thoracic legs pale green. Green
colour phase frequently with pink shading on thorax.

Red colour phase: Head orange-fawn. Prothoracic shield purplish-pink with fawn-
yellow areas laterally. Middorsal band narrow, slate-grey. Subdorsal area consisting
of a dorsal band of grey suffused with pink and a lateral band of purplish-pink; pink
band margined outwardly by a greyish-white line. Supraspiracular area also consisting
of two bands, a dorsal one of purplish-pink and a lateral one of grey suffused with
pink. Spiracular band dull greyish-white. Suprapodal area dull greyish-brown. Mid-
ventral area pale grey. Spiracles with light-brown rims. Setal bases concolorous with
trunk. Thoracic legs fawn.

Head width: 2.68 + 0.12 mm (25 larvae).

Duration of feeding phase of fifth stadium: 7.1 + 1.4 days (68 larvae).

Duration of prepupal phase of fifth stadium: 3.0 + 1.1 days (68 larvae).

Pupa (Figs. 6, 7, 8). Dark orange-brown. Spiracles on a level with general surface
of cuticle; spiracular sclerite weakly projecting. Anterior margins of abdominal seg-
ments 5, 6, 7 each with a narrow band of rather fine pitting. Cremaster consisting of
four elongate setae borne on a short rounded prolongation of the tenth abdominal
segment; one pair of cremaster setae borne ventral or ventro-lateral to the other pair.

Length to posterior margin of fourth abdominal segment: 9.9 + 0.6 mm (18
pupae ).

Acknowledgment

I appreciate the assistance of my associate Mr. E. W. Rockburne, in
measuring larval structures and in drawing the cremaster area of the pupa.

Literature Cited

BEUTENMULLER, W. 1901. Descriptions of three lepidopterous larvae. Jour. New
York Ent. Soc. 9: 90.

Crump, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull.
1135.

Mircu, A. 1868. ‘Twelfth report on the noxious, beneficial and other insects, of the
State of New York. Trans. N. Y. State Agric. Soc. (Part 2) 27: 889-932.

lorspes, W. R. M. 1954. Lepidoptera of New York and neighbouring states. Part 3.
Mem. Cornell Univ. Agric. Exp. Stn. 329.

Gurnir, M. A. 1852. Histoire Naturelle des insectes. Species général des Lepi-
doptéres. Noctuelites. Vol. 2. Paris.

VoLUME 24, NuMBER 4 287

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

Harpwick, D. F. 1967. The life history of Schinia felicitata (Noctuidae). Jour.
Lepid. Soc. 21: 22-26.

Ketuicott, D. S. 1879. An example of protective mimicry. North American Ento-
mologist 1: 30-31.

SAUNDERS, W. 1869. Notes on Alaria florida Guen. Can. Ent. 2: 6-7.

NEW SPECIES OF SYLLEPIS POEY (PYRALIDAE: PYRAUSTINAE),
WITH A KEY TO KNOWN SPECIES

EUGENE MUNROE

Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario

Syllepis semifuneralis Munroe, new species
(Bias, JL, 4)

EXTERNAL CHARACTERS. Head, including palpi, antenna and basal scaling of
proboscis, black; labial palpus narrowly white at base beneath; eye reticulated with
brown. Antenna of male narrowly bipectinate, strongly fasciculate. Body above black.
Body beneath and legs white; front leg shaded with grey dorsally. Posterior part of
abdomen black.

Forewing above Sede. A few white scales on posterior margin basad of middle.
Fringe somewhat brownish.

Hind wing above white. Base narrowly, and apex including fringe, black. A broken
black subterminal line extending from apical patch nearly to anal angle. Some yellow-
ish staining along middle of termen. Posterior part of fringe white.

Wings beneath as above, but hind wing lacking black basal patch and subterminal
line, and without yellowish terminal shading.

Expanse 27 mm. °

MALE GENITALIA. Uncus triangular, about four times as long as basal width,
narrowly rounded at apex, dorsally densely setose in distal third. Gnathos a narrow
band. Subscaphium strap-like. Juxta short, roughly oval. Vinculum prolonged into a
flat, blunt saccus. Valve of moderate width, somewhat expanded distally; costa nar-
rowly inflated; sacculus moderately inflated, with a slender, dorsally directed, slightly
basally curved, spine-like process from its dorsal margin at one-fourth from base to
apex of valve. Penis cylindrical, weakly sclerotized, with a short, blunt cornutus.

FEMALE GENITALIA. Unknown.

TYPES. Holotype male, Bolivia: Dep. Cochabamba, Prov. Chapare,
Alto Palmer, 1100 m. Type No. 11,038, Canadian National Collection.
One male paratype, Peru: R. Inambari, La Oroya, 3100 ft., wet season,
March 1905, G. Ockenden, in British Museum (Natural History).

REMARKS. This species resembles S. religiosa Munroe (1963, p. 704),
but differs in having the forewing almost completely black, and in having
the terminal band of the hind wing obsolete posteriorly.

288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-3. Syllepis spp. 1, S. semifuneralis n.sp., holotype, male; 2, S. triangulifera
n.sp., holotype, female; 3, S. latimarginalis n.sp., holotype, female.

Syllepis triangulifera Munroe, new species
(Figs. 2, 5)

EXTERNAL CHARACTERS. Frons blackish fuscous. Vertex anteriorly fuscous
with a white median patch, posteriorly fuscous. Palpi fuscous; labial palpus fuscous
beneath. Basal scaling of proboscis fuscous. Eye fuscous with blackish strigae. Ocellus
fuscous. Antenna of female filiform, ciliated, fuscous, dorsal scaling shading to grey
distally. Body above black. Head, thorax, and basal part of abdomen beneath and
legs white; front femur, tibia and tarsus infuscated; distal part of abdomen black.

Forewing above black. A broad triangular area based on posterior margin on basal
half of wing white. Fringe black.

Hind wing above white. A minute black basal patch and a black triangular apical
patch of moderate size. Two minute black submarginal dots near anal angle.

Fig. 4. Syllepis semifuneralis n.sp., male genitalia.

VOLUME 24, NuMBER 4 289

Figs. 5, 6. Syllepis spp., female genitalia. 5, S. triangulifera n.sp.; 6, S. latimar-
ginalis n.sp.

290 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Wings beneath as above.

Expanse 30 mm.

MALE GENITALIA. Unknown.

FEMALE GENITALIA. Ovipositor with high, narrow lobes, densely set with
slender setae of varying length. Apophyses slender, nearly straight; anterior apophysis
about twice as long as posterior. Ostium unarmed. Ductus bursae long, slender,
membranous, very finely scobinated, and with an incomplete sclerotized collar near
ostium. Bursa elliptical, finely spinulose, a little more coarsely spinulose on one side.

TYPE. Holotype, female, Peru: R. Inambari, La Oroya, 3100 ft., dry
season, Sept. 1904, G. Ockenden, in British Museum (Natural History).

REMARKS. This species differs from S. semifuneralis in the presence of
a large white area on the forewing and from S. religiosa in the absence of
the terminal band on the posterior half of the hind wing.

Syllepis latimarginalis Munroe, new species
(Figs. 3, 6)

EXTERNAL CHARACTERS. Head, including palpi, eye, ocellus, basal scaling of
proboscis, and basal half of antenna, black. Under side of head and base of palpus
beneath white. Antenna in female annulated, ciliated; distal part brownish. Body
above blackish fuscous. Body beneath and legs white. Dorsal surface of foreleg
greyish fuscous. Tip of abdomen below blackish fuscous.

Forewing broad. Upper surface with costa narrowly and terminal area broadly dark
chocolate brown; costa a little paler just beyond middle. An oblique white band
running antero-distad from basad of middle of posterior margin, ending along and
beyond anterior margin of cell, and with a broadly acute projection into terminal band
behind Cu:. A black line demarcating boundary between white medial and chocolate
terminal bands, and a black line traversing terminal band from apex of projection of
medial band to termen. Fringe blackish fuscous, with pale-buff scaling postapically
and at tornus.

Hind wing above white. A narrow black basal patch. A large, subtriangular, choco-
late-brown apical patch, with irregular black line demarcating its basal boundary.
Traces of a broken black subterminal line on posterior part of wing. Anterior part of
fringe chocolate-brown, posterior part white with lemon-yellow suffusion extending
narrowly onto terminal area.

Wings beneath as above, but all dark areas uniformly blackish fuscous.

Expanse 30 mm.

MALE GENITALIA. Unknown.

FEMALE GENITALIA. Almost as in S. triangulifera, but with ovipositor lobe not
so high, its long setae relatively longer and coarser; anterior apophysis about three
times length of posterior; bursa with fundus rather distinctly globular.

TYPE. Holotype, female, Colombia, West Cordillera, Rio Aguacatal,
2000 m., Fassl, in British Museum (Natural History).

REMARKS. The angled outer edge of the medial band of the forewing
separates this from other species with white ground colour, viz.: S. religi-
osa, S. semifuneralis, and S. triangulifera.

Key to Species of Syllepis

!. Fore- and hind wings above with ground colour straw yellow; distal or apical
areas pink or brown

VoLUuME 24, NUMBER 4 291

Hind wing at least with ground colour white; tip black or dark fuscous. Fore-

mineasimilary coloured or largely infuscated 4
2. Distal part of forewing and apex of hind wing pink_aurora Munroe (1959, p. 359)
Disial parlor sorewing and apex of hind wing brown _... 3
3. Pectinations of male antenna shorter. Continental tropical
A SURGERIES fell hortalis (Walker) (1859, p. 609)
Pectinations of male antenna longer. Antilles marialis Poey (1832, pl. 1)!
4. Forewing above blackish fuscous except for a narrow stripe along posterior
CORR SEEER aE semifuneralis n.sp.
Forewing above with an extensive white patch based on posterior margin 5
5. Distal edge of whitish patch of forewing above nearly straight 6
Distal edge of whitish patch of forewing above forming an acute angle distad
1S REEDS gy ca a a Re er latimarginalis n.sp.
6. Hind wing above with a blackish-fuscous apical patch, followed by one or
two inconspicuous black specks or strigae _________._---_-__-_ triangulifera n.sp.
Hind wing above with a blackish-fuscous terminal band, tapering to a point at
SITE! POPDTACTIT oe ola religiosa Munroe (1963, p. 704)
Acknowledgments

I wish to thank Messrs. P. E. S. Whalley and Michael Shaffer for the
loan of material from the British Museum (Natural History). Two of the
new species were first segregated by Mr. Shaffer. The photographs were
made by Mr. Orrin Hanright, of the BioGraphic Unit, Scientific Informa-
tion Service, Research Branch, Canada Department of Agriculture. Mr.
Douglas Kritsch gave technical assistance in preparing material and
illustrations.

Literature Cited

Munroe, E. 1959. New genera and species of Pyralidae (Lepidoptera). Can. Ent.
91: 359-371.

Munroe, E. 1963. New Neotropical Pyraustinae (Lepidoptera: Pyralidae). Can.
Ent. 95: 704-712.

Pory, F. 1832. Centurie de Lépidopteéres de Vile de Cuba. Paris.

ScHaus, W. 1940. Insects of Porto Rico and the Virgin Islands—Moths of the fam-
ilies Geometridae and Pyralididae. New York Acad. Sci., Scientific Survey of
Porto Rico and the Virgin Islands 12: 291-417.

Waker, F. 1859. List of the specimens of Lepidopterous insects in the collection
of the British Museum. Part 18—Pyralides. London.

1 This couplet follows Schaus (1940: 359), but the marialis group requires revision and will
probably prove to be more complex.

292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NOTES ON TWO RARE MEXICAN ADELPHA AND RELATED
CENTRAL AMERICAN SPECIES (NYMPHALIDAE)

Lee D. MILLER AND JACQUELINE Y. MILLER
Allyn Museum of Entomology, 712 Sarasota Bank Bldg., Sarasota, Florida

Members of the genus Adelpha usually are considered to be tropical
species and usually must be sought in and around the denser moist forests.
The one species that is well known outside the tropics is A. bredowii
(Geyer) which occurs commonly in the southwestern United States and
less abundantly in Mexico and Central America at higher elevations. One
does not usually associate Adelpha with the largely arid Valle de México,
and so we were surprised to find two species rather commonly in the high
country of the state of Hidalgo, México, during January and February,
1969. Neither of these species is well known, and information about them,
as well as comments about other related species should be informative to
other lepidopterists.

Adelpha donysa ( Hewitson )
Figs. 1,2 (¢),9(6¢ genitalia)
yen donysa Hewitson, 1847. Ann. Mag. Nat. Hist., 20: 260 (type-locality:
e€x1co ).
ee roela Boisduval, 1870. Lep. Guatemala, p. 46 (type-locality:
“Mexique’ ).
Adelpha pithys vodena Fruhstorfer, 1915. In A. Seitz, Die Gross-schmetterlinge der
Erde, 5: 524 (type-locality: Guerrero, Mexico) New synonymy.

This species has been rather poorly known since its description. God-
man and Salvin (1884, p. 305) give a rather perfunctory description of
this butterfly, and Fruhstorfer’s (1915, p. 524) description is useless.
Indeed, it appears that Fruhstorfer may have been totally ignorant of the
true nature of donysa and redescribed it under the name of vodena,
though we have not seen the type of the latter name. A. pithys vodena
was described from Guerrero, and the British Museum (Natural History )
contains no material identified as vodena among their huge holdings from
that state, although they do have some typical pithys (Bates) from there.

The present species bears little resemblance, except in a very general
way, to pithys, figures of which are given here (Figs. 3, 4, ; 10, 3 geni-
talia). The two species are in the same general group within the genus,
but clearly distinct. The chief superficial characters separating these but-
terflies are precisely those cited by Fruhstorfer (1915, p. 524) for the
separation of vodena from pithys: the reduced white median bands and
the definitely darker under surface in donysa.

VoLUME 24, NuMBER 4 205

Figs. 1-4. The adelpha pithys group. 1, A. donysa (Hewitson), ¢, upper surface,
México, Hidalgo, 5 mi. NW Zimapan; 2, same specimen, under surface; 3, A. pithys
(Bates), ¢, upper surface, Guatemala, Alta Verapaz, Baleu, Mpio. San Cristdbal
Verapaz; 4, same specimen, under surface.

We encountered donysa in two rather diverse habitats in Hidalgo and
obtained a series of nineteen specimens. The butterfly was most common
in a dry, very “un-Adelpha-like” area five miles northwest of Zimapan,
Hildago, at an elevation of about 2000 m. This area was a dry scrub oak-
juniper association that resembles similar habitats in the southwestern
United States. The “wet” winter landscape is still arid, but the trees are
with leaves, and some herbage is evident. During the summer, however,
a drier place is hard to imagine; the deciduous trees are leafless, and there
is no growth of annuals. The late summer rains are sporadic (in 1967
they never came) so donysa may be univoltine in this area. The second
area was near the village of El Encarnacion, Hidalgo, at elevations be-

294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

tween 2300 and 2400 m, a more inviting place for butterflies in general.
Eumaeus debora Hiibner, a new Callophrys being described by Harry
Clench and Polygonia haroldi C. and R. Felder were among many species
taken there. This locality is something of an oasis in the middle of the arid
mountains of Hidalgo with tall (up to 80-100 feet ) pines and oaks, apple
orchards and rose gardens in the village, comparatively lush under story
vegetation and a permanent stream. Fewer specimens of donysa were
taken there than at the Zimapan locality, however, suggesting that the
area was not arid enough for this unusual Adelpha.

Specimens of donysa were easiest to capture early in the morning, to
about 10:00 AM, when they were sunning and flying near the ground.
This butterfly will sun on the upper side of a leaf or on the ground with
the wings flat and oriented to provide maximum exposure to the sun’s
rays, Obviously a thermoregulatory mechanism (see Clench, 1966, for
details). When not sunning, these butterflies will perch on the uppersides
of leaves in prominent places with the wings upright or slightly open and
are quite aggressive, charging at passing butterflies, birds, and collectors.
The flight at this time is a glide with the wings flat until they reach the
object of their aggression, then they flutter at the object vigorously and
finally break contact and return to their perch. As the day progresses the
butterflies move higher and higher into the canopy, and by mid-afternoon
they may be seen only near the tops of trees. Oaks are preferred by the
adults, and it is probable that the larvae feed on the foliage of these.

The adults show little sexual dimorphism and more closely resemble the
type of roela than that of donysa. It may be that roela is a different, but
not too distinct, subspecies; we have insufficient material at hand to make
such a decision.

Adelpha creton Godman
Figs.5,6(¢4),11(¢ genitalia )
Adelpha creton Godman, 1901. In Godman and Salvin, 1879-1901, Biologia Centrali-
Americana 2: 692 (type-locality: “Jalapa, Mexico” ).

This species was described in the Supplement to the Biologia from a
single female in the Schaus collection. Apparently the type has remained
unique since its description; we have been unable to locate any other
specimens in American or British collections other than the type in the U. S.
National Museum (Type No. 850). In the original description creton was
compared with diocles Godman and Salvin (1884, p. 303), a very similar
insect from the Sierra de Talamanca, Costa Rica, and the Volcd4n de
Chiriqui, Panama, and it seemed reasonable that creton might be a sub-
species of diocles (Figs. 7, 8, é holotype; 12, é genitalia). The genitalia

VOLUME 24, NuMBER 4 295

Figs. 5-8. The Adelpha diocles group. 5, A. creton Godman, 6, upper surface,
Mexico, Hidalgo, vic. El Encarnacion; 2, same specimen, under surface; 3, A. diocles
Godman & Salvin, ¢ holotype, upper surface, Chiriqui, Panama; 4, same specimen,
under surface.

suggest that the two species should be kept separate. It is interesting that
Fruhstorfer (1915) completely ignores both creton and diocles, so it must
be assumed that he had seen specimens of neither. Both species show
pattern convergence toward the south Brazilian abyla complex, but the
two groups are apparently derived from different stocks within Adelpha.

We encountered creton in small numbers in the El Encarnacion area
throughout our stay in Hidalgo. Fifteen specimens were taken, but no
more than two or three in any one day. Many were seen, but they were
high in the oak trees, far out of net range. Occasionally they would glide
down to ground level, alight on dirt or at mud with the wings flat and
nervously move from place to place. These butterflies, to a far greater

296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 9-12. Genitalia of Adelpha. ‘9, A. donysa (Hewitson), ¢ genitalia, slide
M-2112 (Lee D. Miller), México, Hidalgo, 5 mi. NW Zimapan; 10, A. pithys (Bates),
3 genitalia, slide M-2111 (Lee D. Miller), Guatemala, Alta Verapaz, Baleu, Mpio.
San Cristébal Verapaz; 11, A. creton Godman, ¢ genitalia, slide M-2086 (Lee D.
Miller), México, Hidalgo, vic. E] Encarnacion; 12, A. diocles Godman & Salvin, ¢

genitalia, slide TGH 1962-538 (T. G. Howarth, British Mus. (Nat. Hist.), Costa Rica,
Cascajal.

extent than donysa, were very wary and difficult to approach. The inti-
mate association of the adults with the tall oaks of the area suggests that
creton is also a Quercus feeder as a larva.

It seems unlikely that the type of creton actually came from the town of
Jalapa, Veracruz, since that place is in a tropical forest at an elevation of
about 1000 m. It seems more likely that the type of creton came from the
flanks of Mt. Orizaba above Jalapa at an elevation in excess of 2000 m.

VoLUME 24, NuMBER 4 297

This is not the first time that material from the highlands west of Jalapa
in the Schaus collection has been described as from the city. The scarcity
of this species, and for that matter of the Central American diocles, may
be caused by collectors seeking these butterflies in tropical habitats, more
typical for Adelpha, rather than in the high, moist montane valley where
these species occur.

Acknowledgments

We would like to thank Mr. R. I. Vane-Wright of the British Museum
(Natural History) and Mr. William D. Field of the U. S. National Mu-
seum for their help in comparing our material with the types of species in
the respective museums. .

Literature Cited

Ciencu, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47:
1021-1034.

FruustorFEeR, H. 1915. Adelpha. In A. Seitz, Die Gross-schmetterlinge der Erde
5: 510-533.

GopMan, F. D. Anp O. Sayin. 1870-1901. Biologia Centrali-Americana. Insecta.
Lepidoptera-Rhopalocera. London, 2 vols.

THE AFFINITIES OF THE ITHOMIINAE AND THE SATYRINAE
(NYMPHALIDAE)

LAWRENCE FE. GILBERT AND PAuL R. EHRLICH

Department of Biological Sciences, Stanford University, Stanford, California

Fox (1956) called into question the classical association of the nymph-
alid subfamilies Danainae and Ithomiinae, arguing that the Ithomiinae are
actually more closely related to the Satyrinae, especially the glossy-winged
genera such as Pierella. Miller (1968) in his excellent revision of the
Satyrinae quoted Fox’s view and remarked on the presence of a vestige of
vein 3V (3A) as “structural evidence” supporting “the obvious similar
tendency of the ithomiids and the satyrines to have hyaline wings” (p. 15).
As Miller noted, however, this characteristic is also shared by the danaines.
There is, of course, massive morphological evidence clearly showing that
ithomiine adults are more similar to danaines than to satyrines (Ehrlich,
1958, Ehrlich and Ehrlich, 1967). In this paper we present further evi-
dence that Fox’s judgement was in error.

298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

In December, 1969, in the course of a field study of butterfly popula-
tions at the Beebe Tropical Research Station, Simla, Trinidad, W.1., we
used “fedegoso” (dried plants of Heliotropum indicum L. (Boraginaceae )
as an attractant to sample populations of ithomiines for mark-release-
recapture experiments. We were impressed that this plant attracted not
only very large numbers of male ithomiines (Ithomia drymo, Hymenitis
andromica, Tithorea harmonia, Melinaea lilis, Hypothyris euclea, Mecha-
nitis isthmia, and others), but also large numbers of males of the danaine,
Lycorea ceres. No other butterflies were seen to pay the slightest atten-
tion to the fedegoso. A check of the literature reveals that Beebe himself
(1955) had recorded ithomiines and both Lycorea and Danais plexippus
visiting the lure. It is interesting to note here, that on Samoa and other
South Pacific islands, danaines such as Euploea spp. and Danais melissa
(again all males) are attracted by the dead twigs of Tournefortia argentia,
another borage (Buxton, 1926, Poulton, 1936). An additional similarity
among these isolated observations in the Neotropics and in Oceania is that
amatid moths are attracted to these lures in both regions (see also Moss
1947).

Other behavioral traits link the Ithomiinae and Danainae. Both fly in
the sedate manner which is typical of distasteful butterflies, and put on
displays of agility only when attacked. Pierella, on the other hand, is one
of the most rapidly flying satyrines, P. hyalinus displaying behavior char-
acteristic of butterflies with “flash and dazzle” coloration. We have seen
both danaines and ithomiines commonly taking nectar at flowers in the
tropics, behavior rarely encountered in tropical satyrines. On the other
hand, Pierella, Euptychia, and Taygetis all assemble on decaying fruit and
are easily baited with it, but in our experience fruit baited traps have
never attracted ithomiines or danaines. Similar observations on the
absence of danaines on fruit baits have been made in Africa (Rydon,
1964) where danaines were much more prominent in the butterfly fauna.
In Euploea, at least, among the danaines, males are reputed to “assemble”
females (Latter, 1935). It is also strongly suspected that male ithomiines
“assemble” females (Gilbert, 1969). However, this point will remain
uncertain until more is known about the courtship behavior of forest
danaines, ithomiines, and satyrines. Finally, both Lycorea ceres and all of
the ithomiines we have investigated will feign death on capture, often
remaining inert for up to five minutes. We have never observed this
behavior in satyrines.

One might argue that these behavioral similarities are all due to the
mvolvement of the ithomiines and danaines in mimetic complexes, and
imdeed they may be. It should be noted, however, that Heliconius numata

VoLUME 24, NUMBER 4 299

and H. isabella (Nymphalinae), and Dismorphia amphione ( Pieridae,
Dismorphiinae ) are involved in a mimicry complex with Lycorea ceres
and Tithorea, Mechanitis, Melinaea, and Hypothyris around Simla. Al-
though these share flight characteristics with Lycorea, Mechanitis, Meli-
naea, Hypothyris, and Tithorea, they do not visit fedegoso, they do not
feign death, and, at least in the Heliconiini, males do not assemble females.
We consider the morphological and behavioral evidence of the closeness
of adult ithomiines to danaines significant when compared to the super-
ficial similarity in wing transparency of some ithomiines with a few
satyrines.

Similarly, evidence from immature stages indicates close affinity of
ithomiines with danaines, not satyrines. The larvae of the first two always
feed on dicotyledons, the third always on monocotyledons. Both Itho-
miinae and Danainae feed on plants rich in alkaloids, the former almost
exclusively on Solanaceae and the latter primarily on Asclepiadaceae and
Apocynaceae. Furthermore, Tithorea harmonia megara, one of the three
New World ithomiines reported to feed on non-solanaceous plants, feeds
in Trinidad on Echites (= Mesechites trifida Jacq., Apocynaceae; Guppy
1894, 1904). In Costa Rica, Fountaine (1911) reared Tithorea “pinthias”
(= T. tarricina duenna Bates; see Fox, 1968) on a plant which from its
description is probably Echites or another Apocynaceous vine. Guppy
(1894) also reports finding Sais (= Aeria) eurymedia on Echites in
Trinidad.

Morphologically, larvae of the danaine Lycorea and the ithomiine
Tithorea are strikingly similar (descriptions in Guppy, 1894, 1904 and
Fountaine, 1911 and illustrations in Guppy, 1904), and when disturbed
have the same behavior of nervously twitching the long flexible protuber-
ances which arise from the first segment behind the head. Guppy (1894)
describes A. eurymedia larvae as almost identical to those of Tithorea
except for their much smaller size at pupation. The larvae of such itho-
miine genera as Dircenna, Hypothyris, Ithomia, Godyris, and Mechanitis
do not superficially bear such strong resemblance to the danaines, nor to
one another for that matter (Gilbert, personal observation). In any case,
ithomiine larvae do not seem suggestive of the satyrines.

Miller (1884) in his classic work “Sudamerikanische Nymphaliden
Raupen” pointed out two characteristics in common among danaines and
ithomiines. One is primary bristle development around larval bristle la.
The second major characteristic suggesting close relationship is the
rigidity of the pupae. The only two major differences between these
groups according to Miiller are, firstly, that danaines possess and itho-
miines lack the “Scherndornen” (long flexible protuberances), and sec-

300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ondly that danaines feed on Asclepiadaceae whereas ithomiines feed on
Solanaceae. Unfortunately, Muller apparently never worked on the life
history of Tithorea, which represents a major exception to both of his
criteria for separating these two groups.

Acknowledgments

We would like to thank Michael C. Singer for advice in translating the
Miller paper.

Literature Cited

BEEBE, W. 1955. Two little-known selective insect attractants. Zoologica 40: 27-32.

Buxton, P. A. 1926. Euploea spp. frequenting dead twigs of Tournefortia Argentea
in Samoa and Tonga. Proc. Ent. Soc. Lond. 1: 35-37.

Enruicu, P. R. 1958. The comparative morphology, phylogeny and higher classifi-
cation of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull.
39: 305-370.

Euruicu, P. R. AnD ANNE H. Enruicu. 1967. The phenetic relationships of the
butterflies. I. Adult taxonomy and the nonspecificity hypothesis. Systematic
Zoology 16: 301-317.

FounTAINE, M. E. 1911. Five months butterfly collecting in Costa Rica in the
summer of 1911. Entomologist 46: 189-194.

Fox, R. M. 1956. A Monograph of the Ithomiidae (Lepidoptera) Part I. Bull.
Amer. Mus. Nat. Hist. 111(1).

Fox, R. M. 1968. Ithomiidae (Lepidoptera: Nymphaloidea) of Central America.
Trans. Amer. Ent. Soe. 94: 155-208.

GILBERT, L. E. 1969. Some aspects of the ecology and community structure of itho-
mid butterflies in Costa Rica—Mimeo Res. Rep. Adv. Pop. Bio. Organization for
Tropical Studies, Costa Rica, pp. 69-93.

LatrerR, O. H. Aanp ELTrincHaM, H. 1935. The epigamic behavior of Euploea
(Crastia) core asela (Moore) (Lepidoptera Danainae). Proc. Roy. Soc. (B)
117: 470-482.

Miutuer, LEE D. 1968. The higher classification, phylogeny, and zoogeography of
the Satyridae (Lepidoptera). Mem. Amer. Ent. Soc. 24.

Moss, A.M. 1947. Notes on the Syntomidae of Para with special reference to wasp
mimicry and fedegoso, Heliotropium indicum (Boraginaceae) as an attractant.
Entomologist 80: 30-35.

Mutter, W. 1886. Sudamerikanische Nymphaliden Raupen. Zool. Jahrb. I.

Poutton, E. B. 1936. Euploeine butterfly feeding at the same broken surface of a
Tournefortia branch and some days later on the withered leaves of the same
branch: Tulagi, Solomon Islands 1936. Proc. R. Ent. Soc. Lond. (A) 11: 94-96.

VoLUME 24, NuMBER 4 301

A DAY-FLYING MOTH (PERICOPIDAE) NEW TO TEXAS
AND THE UNITED STATES

Roy O. KENDALL
135 Vaughan Place, San Antonio, Texas

Phaloesia saucia Walker

Phaloesia saucia Walker, 1854. List of specimens of lepidopterous insects in the col-
lection of the British Museum, Part 2, p. 359 (genus and species described from
4 specimens, two from Venezuela, one from Nicaragua, and one from unknown
locality ); Butler, 1874, Trans. Ent. Soc. London (1874) p. 54; Druce, 1884, Biol.
Centr.-Amer., Insecta Lep., Het. 1: 105, t. 11, f. 2; ibid., 2: 384 (1890); Kirby,
feo syne Cat. ep, Het! 1: 89) n. 1. Dyar, 1911, Proc. Ent. Soc. Wash. 13:
230; Dyar, 1914, Ins. Inscit. Menstr. 2: 63; Hering, 1925, in Seitz, Die Gross-
Schmetterlinge der Erde 6: 448; Bryk, 1931, in Strand, Lepid. Cat. Part 45, p. 40.

Cocastra gentilis Boisduval, 1870, Consid. Lep. Guat. p. 88 (synonymy ).

Phaloesia fulvicollis Butler, 1875, Ann. Mag. Nat. Hist. (4) 16: 171 (synonymy ).

Phaloesia chalybdea Butler, 1875, Ann. Mag. Nat. Hist. (4) 16: 171 (synonymy ).

Phaloesia veneszuelae Butler, 1875, Ann. Mag. Nat. Hist. (4) 16: 171 (synonymy).

The last four names were praced in synonymy bv Druce (1884) to which Butler
agreed.

McDunnough (1938) does not list this species as being found in the
United States, even though examples had been collected at Brownsville,
Cameron County, Texas 28 years earlier. Because few collectors may be
familiar with this species, a male and female are illustrated. There is con-
siderable size variation in each sex, possibly due to availability or scarcity
of larval foodplant. Of the nine examples collected by the writer, the wing
expanse of males ranged from 1% to 1’ inches, and of females from 1% to
2 inches.

Dyar described the larva, cocoon, and pupa but did not give its larval
foodplant. This species has been collected at or near Brownsville, Texas
in April, May, August, October, and November. Collection dates would
indicate there are at least three broods in extreme southern Texas where it
has been taken by various collectors over the past 60 years. It is therefore
considered resident.

Specimens observed and collected by the writer have been in thick
brush or on the edge thereof, making it difficult to catch except when
feeding on blossoms. This dense and often thorny habitat probably
accounts for the tattered condition of some specimens.

The earliest known records for the United States are three examples in
the U. S. National Museum. A female bears the labels “Brownsville, Tex.,
Nov. 30, 1910”; “Sweeping weeds, STC., Nov. 20, *10”; and “Phaloesia

302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

2

Figs, 1,2. Phaloesia saucia Walker, dorsal view; Brownsville, Tex., 3.1V.1957
(R. O. Kendall). 1, Male (expanding 1.5”); 2, female (expanding 2.0”).

saucia’. The other two specimens, a 6 and ? collected in copula, Browns-
ville, Texas April 7, 1929, F. H. Benjamin ( 6 very battered and worn).

The writer has collected nine examples at or near Brownsville. In April
1957 it was found fairly common, but not realizing it to be a good catch,
only a few examples were taken. Specific dates are: 2 April 1957 (14,
229 ),3 April 1957 (26 6,12), 7 November 1969 (2 fresh 2 2 feeding on
blossoms of Eupatorium odoratum L.), 9 November 1969 one sight record,
and 10 November 1969 (12). Other examples were seen on 7 November
but the habitat made it impossible to catch them.

Jack E. Lipes (1962) found it at Brownsville in early May and again in
August. In personal correspondence, he informed the writer of taking it in
flight 7 August 1961.

Michael A. Rickard (personal communication) collected it at Browns-
ville 31 May 1968 (12), 25 October 1969 (14), and 8 November 1969

are
| | }

VOLUME 24, NuMBER 4 303

Acknowledgments

I wish to thank Dr. E. L. Todd, Systematic Entomology Laboratory,
U.S. Dept. Agriculture, at the U. S. National Museum of Natural History
for providing data on specimens in the U.S.N.M.., for the reference cita-
tions, and for reviewing the manuscript. To André Blanchard, a very
good friend and companion on numerous field trips throughout Texas, I’m
grateful for photographing the specimens illustrated.

Literature Cited

Lives, Jack E.. 1962. Season Summary (1961). News Lepid. Soc. No. 3.
McDunnouwcu, J. 1938. Check list of the Lepidoptera of Canada and the United
States of America—Part I, Macrolepidoptera. Mem. S. Calif. Acad. Sci. 1.

NOTES ON THE USE OF NET-TRAPS AT PALAWAN, PHILIPPINES

In past issues of this journal, I came across two articles dealing with the preparation
and use of net-traps provided with baits to catch certain species of high flying forest
butterflies, especially Charaxes and other rarely seen nymphalids. Presently there
seems no record to show that such type of traps have been tried in the Philippines.

During the writer’s fourth expedition to Palawan for the further study of the biology
of the rare Trogonoptera trojana and other papilionids of the forest, we took along
huge mosquito nets. These we intended to use as a hatchery for forest butterflies, and
for closeting immatures right on their host plants. With the latter, we were unsuccess-
ful, but we found other uses for these large nets.

We selected river bends or spots where two creeks converge, and also promising
creeksides, and near these we hung the mosquito nets, using nearby trees or poles to
hold them. The lower edges of the nets were about five inches above the ground and
held firmly with stone anchors. Between July and October, the mountain creeks and
streams at Palawan have a heavy traffic of roving pierids, certain nymphalids, and
some papilionids. Three nets were constantly in use. Two of medium sizes were
installed permanently by the creekside, and a very large one was set at several places.

In an issue of this journal in 1958, the author discussed his experience in the use of
baits to attract butterflies. This time, because of the duration of our stay in the forest,
an opportunity to try baiting butterflies once more presented itself. At first we used
overripe fruits of the ates (Anonas squamosa L.) which we placed on the balcony of
our hut. This was at an altitude of about one thousand feet in a densely forested
valley. Early each night, some three species of Underwing Moths often came to the
bait. These moths showed a high preference for this fruit over several others like
guavas, jackfruit, and papaya.

Beginning in the month of August when it was less windy than in July, we set the
nets. In its early phase, our baiting attempt concentrated on the use of overripe fruits,
mashed pineapple jam with rum and tid-bits from our table. In this case, we were able
to collect a fine series of forest butterflies such as a female Zeuxidia, euthalids, Adolias,
Pantoporia, Precis, Phalanta, Eulepis, Charaxes, Libythea, and some moths. Some of
these visitors came at twilight, but most of them, by day. In a small coppice near a
large creek, thousands of flies of several species swarmed into the trap.

304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

A most astounding result was noted when, for a joke, someone placed human feces
inside the trap. Since the initial result seemed to indicate that certain butterflies,
especially nymphalids, showed an unusual liking for this bait, it was decided to try
alternating overripe fruits and human excreta, and then concentrate on the one that
proved more fruitful. In a short time, it was found that human feces placed early in
the trap would attract more visitors near noon on hot days than anything else. This
type of bait therefore was employed for about two months, and brought a rich harvest
of nearly five hundred and fifty butterflies, some of which are hard-to-get forest
nymphalids.

A list of species, taken in traps baited with human feces, follows: 220 Eulepis
delphis niveus Roths., 105 Vindula arsinoe palawanica Fruh., 35 Charaxes of some
three species, Graphium jason, Papilio lowii, P. palawanicus, Phalanta, Cupha, Precis,
Libythea, Appias nero palawanica Staud., A. albina agatha Stgr., Symbrenthia,
Huphina, Cyrestis maenalis obscurior Stgr., Discophora tullia semperi Mre., Eurema,
and several species of skippers. Possibly, if the nets were hung at various elevations,
or placed on other terrain with different vegetational associations, the catch would also
include a good number of forest butterflies that are seldom seen near the ground.
Smaller nets such as those described in this journal might also be effective if hung in
trees where species adapted to the upper layers (emergent canopy ) are likely to catch
the scent of the bait used.

Along the creeksides where the nets were located, hundreds of lively pierids, such as
Appias nero and A. albina, passed daily up- or downstream. While the white Appias
were frequently attracted to the bait, the red one was seldom noted to come near it.
On sunny days, the smallish Palawan Vindula and Eriboea (Eulepis) were active
mostly between ten in the morning and a little past noon. The pierids had a similar
period of flight but they were still active at three in the afternoon on sunny days.
Most of the trapped Charaxes were males. Only rarely were females taken. Of the
mountain swallowtails, it was interesting to note that two, Papilio palawanicus and
lowii, were frequent visitors to the bait. There were only two or three records of the
P. polytes being attracted to human feces.

The accidental use of large net-traps at Palawan, together with the equally un-
planned introduction of human feces as bait, has brought to our collecting group at the
University of San Carlos a new technique for collecting at places far from human
habitations. The large size of the net easily enabled the collector to enter the trap and
pick the fluttering prisoners at the net’s corners where they usually retire when tired.
We also noted that when we were away for two or three days, we would be greeted
by an accumulated catch, unless a passing dog had messed up everything.

One handicap with setting nets on the ground or close to it, is their molestation by
passing dogs and night prowlers such as civet cats, wild pigs, and bearcats. In this
case, if very large nets were employed, it may fare better if bamboo poles are used as
weights of the edges of the net. The corners could be anchored to the ground with
pegs or stones. The firm edges would discourage animals from crawling into the bait.

This note does not present any particular design and size for the nets we employed,
because ready-finished mosquito nets were used. It would be best to experiment with
different sizes for use on various types of terrain.

JuLian N. Jumaton, Department of Biology, University of San Carlos, Cebu City,
Philippines. |

VoLUME 24, NuMBER 4 305

SOME BUTTERFLIES OF MADISON COUNTY, ILLINOIS

In the years since I’ve been collecting butterflies I have made numerous trips to my
inlaws’ country home about 114 miles west of the crossroads community of Prairie-
town, Madison County, Illinois. This is about 30 miles northeast of downtown St.
Louis, Missouri. When time, weather, and season allowed I have collected butterflies.
Recently I was reading some old journals and the President's address by Munroe
(1960, J. Lepid. Soc. 14: 1-4), along with some urgings from my wife, prompted me
to write this record of my captures.

The area which was collected consists of about eight acres of mowed grass and
about 14 acres of pasture land which was abandoned as such about five years ago and
allowed to grow wild. There is a small pond in the mowed area around which the
grass is allowed to grow to its natural height. Along the highway the grass was more
or less of natural height also.

Slightly more than 950 specimens representing 56 species were captured on the 67
days that at least some collecting was done in the area. The majority of the collecting
was done around the three holidays, Memorial Day, the 4th of July, and Labor Day.
This by no means exhausts the different butterflies to be caught in this area for in
1969 alone I added six to my list. In addition there are nine other species I have
caught in one year only.

The 56 species so far caught can normally be found in the area at one time or
another according to Klots (1951, A field guide to the butterflies) and are listed
below in the order of Dos Passos (1964, Synonymic list of the Neararctic Rhopalocera):

Amblyscirtes samoset (Scudder )
A. vialis (Edwards ) Eurema lisa Boisduval & Le Conte
Euphyes vestris ( Boisduval ) Strymon acadica (Edwards)
Poanes zabulon ( Boisduval & Le Conte ) S) edivardsit (Grote & Robinson)
Atalopedes campestris ( Boisduval )
Pompeius verna (Edwards )
Wallengrenia otho (Smith)

Polites coras (Cramer )

P. themistocles ( Latreille )

P. origines ( Fabricius )
Ancyloxypha numitor ( Fabricius )
Pholisora catullus ( Fabricius )

Phoebis sennae ( Linnaeus )

Lycaena thoe Guerin-Meneville
L.. xanthoides ( Boisduval )

L. phlaeas ( Linnaeus )

Everes comyntas ( Godart )
Celastrina argiolus ( Linnaeus )
Libytheana bachmanii ( Kirtland )

Asterocampa celtis (Boisduval & Le Conte)

Pyrgus communis (Grote )
Staphylus mazans ( Reakirt )
Thorybes bathyllus (Smith)
T. pylades (Scudder )

T. confusis Bell

Achalarus lyciades ( Geyer )
Epargyreus clarus (Cramer )

Battus philenor (Linnaeus )
Papilio polyxenes Fabricius
P. cresphontes Cramer

P. glaucus Linnaeus

P. troilus Linnaeus

Pieris protodice Boisduval & Le Conte

P. rapae ( Linnaeus )
Colias eurytheme Boisduval
C. philodice Godart

A. clyton ( Boisduval & Le Conte )
Limenitis astyanax (Fabricius )

L. archippus (Cramer )

Vanessa atalanta ( Linnaeus )

V. virginiensis ( Drury )

V. cardui (Linnaeus )

Junonia coenia ( Hubner )
Polygonia interrogationis ( Fabricius )
P. comma ( Harris )

Phyciodes tharos ( Drury )
Melitaea nycteis Doubleday
Speyeria cybele (Fabricius )
Euptoieta claudia (Cramer )
Danaus plexippus (Linnaeus )

Lethe portlandia ( Fabricius )
Euptychia cymela (Cramer )
Cercyonis pegala (Fabricius )

Cart R. Cusuine, 2146 34th St., Los Alamos, New Mexico.

306 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

NOTES ON A BRIEF COLLECTING TRIP TO NEGROS, PHILIPPINES

A Yuletide collecting trip to the hydro-electric plant site on the island of Negros,
Philippines, occasioned a four-day observation of mountain satyrids and a few
nymphalids under continuously bad weather. The water power plant is located on a
well-forested mountain with elevations from one thousand to three thousand feet.
Deep gorges and densely forested sharp slopes are characteristic of this area, but there
are good trails into the forest running in many directions from the main road. These
trails have been found rich in many species of nymphalids and forest satyrids.

Several days prior to our visit, the roadsides were cleared of tall ferns and grasses.
One species of grass, with stiff stems as large as pencils, secreted a sap which had fer-
mented after a few days. This was found to attract nearly ten species of satyrs,
several of Neptis as well as three amathusiids. So, despite the rains and fogs during
our four days of collecting, we were able to pick a good number of these visitors to
the stumps of the grass. I was especially interested in the two species of Ptychandra
and three of mountain amathusiids, a Discophora, an Amathusia, and a Clerome.
Since this species of grass is common in the area and also on several islands, the dis-
covery will help in future expeditions to increase the catch of these types of forest
butterflies. I have not as yet determined the number of days before the exuded sap
begins attracting the insect visitors.

We had no chance to observe the “bait” during sunny days because we never saw
the sun during our stay. Our cold nights were spent around the posts of mercury-
vapor lamps to collect the numerous sphinx moths and many smaller species, as well
as some three species of Antheraea, one of a long-tailed Luna Moth, and an interesting
mountain species of Attacus. The use of mercury-vapor lamps at logging and mining
areas in mountainous areas makes such places rich collecting grounds for moths of all
kinds, including several huge beetles. Because of the findings at Negros, a formal
expedition to the place is being planned for the coming hot season in the Philippines,
to determine if the dry season is a better collecting time in the rain forest of Negros
island. Negros has a seven-thousand foot volcano named Kanla-on with a dense forest
surrounding it. The lowlands are well adapted to sugar-cane, the main crop of this
province.

Juuran N. Jumaton, Dept. of Biology, University of San Carlos, Cebu City, Philip-
pines.

THE FEEDING HABITS OF OENEIS JUTTA REDUCTA

A recent paper by Masters and Sorensen (1969, J. Lep. Soc. 23: 155-161) dis-
cussed the habits of several species of Oeneis. They mentioned, quoting from Eff in
Brown (1957, Colorado Butterflies), that Oeneis jutta reducta McDunnough “. . . in
spite of the multitude of flowers nearby never visited any.” The conclusion was that this
species is not a flower feeder. I have collected this insect at six locations in four coun-
ties in Wyoming. In two of these areas, it was found to be an avid flower feeder. The
habitats vary from exceedingly dry lodgepole pine forests, far removed from moist areas,
to the borders of willow bogs such as described by Masters and Sorensen.

The first specimens taken on flowers were collected along the Doubletop Mountain
Trail, Bridger Wilderness Area, Sublette Co., Wyoming on 22 July, 1969. I took four
specimens and saw others on flowers in a boggy meadow at the forest’s edge. On the
same date, other members of the collecting party (D. R. Groothuis, J. D. Eff, and P. J.
Conway) collected additional flower-feeding specimens. The insects exhibited a
preference for yellow flowers although some were seen feeding upon white ones. The

VoLUuME 24, NuMBER 4 307

butterflies, while feeding, remained quite motionless with wings tightly closed. They
seemed quite unwary and were easily approached, which is not the usual case with
this species in heavily wooded areas.

Additional specimens of O. jutta reducta were taken on yellow flowers along a dirt
road in Albany Co., Wyoming on 1 August, 1969. They were found at the edge of a
very dry lodgepole pine stand.

Other specimens collected during 1969 in Sublette, Fremont, and Sheridan Cos.,
Wyoming were found in rather heavily wooded areas flying about rocks and deadfall
as described in Brown’s book.

From these observations, it must be concluded that O. jutta reducta does feed upon
flowers in some instances. Both sexes were involved. Flower feeding was observed
during the hours from 11:00 AM to 2:00 PM (MDT).

CuirrorD D. Ferris, College of Engineering, University of Wyoming, Laramie.

BOOK REVIEW

ANNOTATIONES RHOPALOCEROLOGICAE 1970—Charles F. Cowan, Clunbury Press, Berk-
hamsted, Herts, England, 70 pp., 15 April 1970. $3.00 US.

“This is the second and final instalment of an addenda and corrigenda to the late
Francis Hemming’s “Generic Names of the Butterflies and their Type-Species’ . . .” is
the opening sentence of this most interesting booklet. Col. Cowan’s two contributions
to the stability of nomenclature must be studied by every nomenclator and every active
taxonomist. Here is a synopsis of the current part:

A. An addenda of generic names published before 1965 and omitted from Hem-
ming’s work. Thirty-two names, including eleven for American Lycaenidae by Clench
and Poladryas Bauer, are discussed. Seventy names omitted from the index of
Hemming’s book are listed.

B. Comments on dates of publication of works by 17 authors, including several
written by S. H. Scudder.

C. Comments on the changes made in 1964 relative to Articles 11 and 31 of the
International Code. These affect forty names including several genera appearing in
the Americas.

D. Comments on several cases pending before the I.C.Z.N. involving butterfly
genera.

E. An analysis of emended names for genera and comment upon the justification or
lack of justification for each case. Several of the forty-seven names touched upon apply
to American butterflies.

F. This particular section is an index to all names commented upon by Cowan. The
alphabetic list of emendations to be made to Hemming’s “Generic Names . . .” is
extremely useful.

G. Here are listed alphabetically: incorrect subsequent spellings and unjustified
emendations, unadopted synonyms, and ghost names.

Copies of this valuable little booklet may be purchased from the author at Little
Gaddesden House, Berkhamsted, Herts, England.

F. Martin Brown, Fountain Valley Rural Station, Colorado Springs, Colorado.

308 JouURNAL OF THE LEPIDOPTERISTS SOCIETY

INDEX TO VOLUME 24

Subject Index californica Wright, 226
Callithomia, 23

annual meeting, 73 Callophrys, 37, 228
bait trapping, 303 Cameraria, 86
behaviour, 72, 117, 189, 190, 226, 270, cardui Linn., 157

271, 306 Catocala, 105
book reviews, 228, 307 Cautethia, 267
checklists, 26 Chlosyne, 135, 226
collecting notes, 306 Cithearias, 17
collection evaluation, 51 elenchi Miller, 122
development, 209 clymene Brown, 271

distribution, 143, 202, 203, 212, 217, 226. Ccenonympha, 202
297, 228, 233, 240, 244, 249, 266, 292, Colias, 117

301 comstocki Henne, 228
life history, 47, 54, 135, 151, 227, 234, Coryphista, 220

282 Crambidia, 227
light trapping, 19 creola Skinner, 145
migration, 157 cresphontes Cram., 239
nomenclature, 38, 76, 143 creton Godman, 294
photography, 61 daira daira Godart, 240
regional lists, 1, 42, 305 daira palmira Poey, 240
seasonal abundance, 105 Dasypyga, 253
technique, 81 dion Edw., 243
transferal of collection, 208 donysa Hewitson, 292

variation, 77, 125, 176, 224, 229, 239 dorantes dorantes Stoll, 241, 245
Ecpantheria, 227
Name Index Euchloe, 226
(New names in boldface ) Eumaeus, 36

Euphyes, 243
aaroni howardi Skin., 243 ae 240
Actias, 209

: é eurytheme Bdv., 117
actorion actorion Linn., 18

Eutricopis, 151
Adelpha, 292 exilis yucateca Clench, 4
Allosmaitia, 60

i Wilk., 234
allottei Rothschild, 88 SAS UU ROIS 5

florida Gn., 282
ancellata Hulst, 54 Formicidae, 190
ancillaris, 266

fotis schryveri Cross, 256
andromeda Fabr., 17 foxd Rrcet 23
antimachus Dru., 224 fragariae iG. Edw., 189
Arctiidae, 227, 271 freemani Miller, 120
aristolochiae Fabr., 19

Gelechiidae, 6
aspasia Murayama, 213 Cepmaniias. BL 220)
aspasia coreensis Murayama, 215 Gonepteryx 913. 218
aspasia iwateana Murayama, 214 fivien 1 sae * 9929
aspasia kansuensis Murayama, 215 Bee aaa ae

Ses eae saa Gracillaridae, 86, 272
astyoche Erichson, 17 ;
Atlides, 37 Graphium, 176
ausonides mayi Chermock & Chermock, Haetera, 16
226 haferniki Freeman, 247
boreale Lafontaine, 83 Haploa, 271
borealis Lafontaine, 83 Harkenclenus, 36
Bia, 18 Hesperiidae, 2, 45, 47, 120, 247, 266

Brephidium, 4 Heterosmaitia, 36

VoLUME 24, NuMBER 4

hyalinus hyalinus Gmelin, 17

Incisalia, 256

inornata Edw., 202

Ithomiidae, 22

jutta reducta McD., 306

Keiferia, 6

kendalli Blanchard, 249

kenora Free., 280

lacinia Geyer, 135

latimarginalis Munroe, 290

Lerema, 266

Lethe, 143

Limentis, 270.

Lithocolletis, 273

luna Linn., 209

Lycaenidae, 2, 3, 26, 44, 56, 59, 83, 190,
256

macrocarpae Free., 86

marcellus Cram., 176

meadi Pack., 220

Megathymidae, 208

Mestra, 203

napi oleracea Harr., 77

neumoegeni Skin., 226

nexilis Morr., 151

nipigon Free., 276

Noctuidae, 3, 105, 151, 234, 282

noctuiformis bredini Cary, 267

Nymphalidae, 10, 25, 43, 135, 157, 203,
226m 20s 2925, 297

Ocaria Clench, 56, 60

ocrisia Hewitson, 60

Oeneis, 306

ontario Free., 273

Ornithoptera, 88

palegon Stoll, 60

Papilio, 224, 229, 239

Papilionidae, 19, 45, 176, 229, 233, 239

Pericopidae, 301

Perisama, 10

Phaloesia, 301

Phoebis, 118

piera piera Linn., 16

Pierella, 17

ienGacr4o. vo lil? 125, 93, 294 296

Richisw (125,224 DA5

pion Godman & Salvin, 60

Piruna, 247

Poanes, 243

Polydorus, 19

populiella Cham., 279

proteus Linn., 47

protodice protodice Bdv. & LeC., 245

pura B. & McD., 227

Pyralidae, 249, 287

309

Pyrrhia, 234

Ramosia, 189

rapae Linn., 125, 224
rhamni tianshanica Nekrutenko, 218
Riodinidae, 13

rusea Masters, 10
rusposoria Povolny, 6
salicivorella Braun, 279
salmocolor Blanchard, 253
saucia Wlk., 301

Satynidaes iy 15) 43" 1435 902
Satyrium, 36, 83

Schinia, 282

Scopula, 54

scribonia Stoll., 227
scudderella Frey & Boll, 278
semifuneralis Munroe, 287
sennae eubule Linn., 118
Sphingidae, 267

Strymon, 83

Syllepis, 287

Thereus, 58, 60
tremuloidiella Braun, 275
triangulifera Munroe, 288
Wrlamusy: 4740241, 24'5
Vanessa, 157

Vinius, 120

Virga, 122

Zamagiria, 249

Author Index

Berube, 220
Blanchard, 249
Brou, 239

Brown, 307

Bush, 135
Callaghan, 228
Carcasson, 72

Cary, 267

Clarke, 229

Clench, 3, 56, 117, 240
Concello, 227
Covell, 51
Cunningham, 271
Cushing, 305
Diakonoff, 70

Doyle, 212
Drummond, 135
Ehrlich, 224, 297
Bichlin, 271

Emmel, 135

Emsley, 25

Ferris, 202, 256, 306
Freeman, H. A., 68, 247

310

Freeman, T. N., 86, 272
Gage, 270

Gilbert, 297

Greene, 47

Hardwick, 151, 228, 234, 282
Hessel, 105

Howe, 217

Irwin, 143

Jae, 233

Jumalon, 303, 306
Kendall, 59, 266, 301
Kolyer, 61, 125
Lafontaine, 83

Leech, 189

Malicky, 190

Masters, 10, 15, 22, 203
Mather, 176

McGuffin, 54

Miller, J. Y., 244, 292
Miller, L. D., 13, 120, 244, 292
Moiz, 19

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Muller, 1
Munroe, 287
Munshi, 19
Nekrutenko, 213, 218
Nielsen, 42
Oliver, 77
dos Passos, 26
Povolny, 6
Priestaf, 226
Rindge, 208
Riotte, 227
Rozman, 81
Sargent, 105
Schmid, 88
Shapiro, 224
Sheppard, 229
Stanford, 256
Steyskal, 38
Tasker, 226
Williams, 157
Wright, 209

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NOTICE TO CONTRIBUTORS

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of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
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Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
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Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA
by Cyrit F. pos Passos
Price, postpaid: Society members—$4.50, others—$6.00; uncut,
unbound signatures available for interleaving and private binding,

same prices; hard cover bound, add $1.50. Revised lists of the
Melitaeinae and Lycaenidae will be distributed to purchasers free.

ALLEN PRESS, INC. ra pateee LAWRENCE, KANSAS
usr

CONTENTS

Blanchard, A. Observations on some Phycitinae (Pyralidae) of Texas with

descriptions of two new species 2 .0i 1 en ee 249-255
Brou, V. A. A melanic aberration of Papilio cresphontes (Papilionidae ) 239
Brown, F. M. Book Review: Annotationes Rhopalocerologicae clin

Cary, (Mrs.) C. R. A new sphinx moth from the West Indies (Sphingidae) 267-270

Clarke, C. A. and P. M. Sheppard. Is Papilio gothica a good species -__.. 229-233
Clench, H. K. New or unusual butterfly records from Florida —_...-_ 240-244
Cushing, C. R. Some butterflies of Madison Co., Illinois a 305
Eichlin, T. D. and H. B. Cunningham. Larvae of Haploa clymene ( Are-

tiidae) hibernating on concrete bridges _.-.--- pag
Ferris, C. D. The feeding habits of Oeneis jutta reducta (Satyridae) 306-307
Ferris, C. D. and R. E. Stanford. Incisalia fotis schryveri (Lycaenidae):

bionomic notes and ‘Jife history. 256-266
Freeman, H. A. A new species of Piruna from Texas (Hesperiidae) ___ 247-249
Freeman, T. N. Canadian species of Lithocolletis feeding on Salix and

Populus: (Gracillariidae), 272-281
Gage, E. V. Record of a naturally occurring Limenitis hybrid (Nymphal-

Bebe) AS ae Uo ac 270
Gilbert, L. E. and P. R. Ehrlich. The affinities of the Ithomiinae and

Satyrimae 00 S08 Cie Oe MIO ESE INN, CAT hep 297-300
Hardwick, D. F. The life history of Pyrrhia experimens (Noctuidae) __. 234—239
Hardwick, D. F. The life history of Schinia florida (Noctuidae) _________ 282-287
Jae, R. J. A new record for North America of a swallowtail butterfly (Pa-

Piliomidae ) FNS STO OS 233
Jumalon, J. N. Notes on a brief collecting trip to Negros, Philippines 306
Jumalon, J. N. Notes on the use of net-traps at Palawan, Philippines _____ 303-304
Kendall, R. O. A day-flying moth (Pericopidae) new to Texas and the

United. States ou Moos Waa Su IST Cn nares ee 301-303
Kendall, R. O. Lerema ancillaris (Hesperiidae) new to Texas and the

United States tse PS UN Ue rr 266
Miller, L. D. and J. Y. Miller. Pieris protodice and Urbanus dorantes in

southern Florida) 0000) Ni) Vari SN Onan ei ae ne 244-247
Miller, L. D. and J. Y. Miller. Notes on two more Mexican Adelpha and re- :

lated Central American species (Nymphalidae) _ 292-297

Munroe, E. New species of Syllepis Poey (Pyralidae: Pyraustinae), with
a key to known species 200/00 OM i TT aT 287-291

Index to Volume 24

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Volume 25 1971 Veg | Number 1

JOURNAL

of the

LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

23 February 1971

THE LEPIDOPTERISTS’ SOCIETY

EprroRIAL COMMITTEE

D. F. Harpwick, Editor of the Journal
C. V. Covet, Editor of the News
S. A. HesseL, Manager of the Memoirs

ExEcuTIvE Councr (1970)

E. B. Forp (Oxford, England), President

C. L. Remincton (New Haven, Conn.), President-elect
L. P. Brower (Amherst, Mass.), Ist Vice President

L. M. Martin (Prescott, Ariz.), Vice President

J. W. TrtpEN (San Jose, Calif.), Vice President

S. S. Niconay (Virginia Beach, Va.), Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

Members at large (three year term): W. C. McGurrin (Ottawa, Ont.) 1971
J. F. G. Cuarxe (Washington, D.C.) 1970 Y. Nexrurenxo (Kiev, U.S.S.R.) 1971
H. K. Ciencu (Pittsburgh, Penna.) 1970 B. Maruer (Clinton, Miss.) 1972

B. Wricut (Halifax, N.S.) 1970 M. Ocata (Osaka, Japan) 1972

A. E. Brower (Augusta, Me.) 1971 E. C. Wexuinc (Merida, Mexico) 1972

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
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JOURNAL OF

Tue LEPIDOPTERISTS’ SOCIETY

Volume 25 1971 Number 1

THE LIFE HISTORY OF HELIOTHIS
OREGONICA (NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute,
Canada Department of Agriculture, Ottawa, Ontario

Heliothis oregonica (Hy. Edwards, 1875) is widespread in montane
western North America, occurring from southern British Columbia south-
ward in the Sierra Nevada-Cascade Axis to the Lake Tahoe area of Cali-
fornia and in the Rocky Mountain System to southern Colorado. In west-
ern America, its altitudinal range is rather wide: in southern British
Columbia, for example, it occurs at altitudes between 1500 and 6000 feet.
Until recently the species was considered to be confined to the West;
however, two specimens were taken in 1956 at Mistassini Post in central
Quebec. A single additional specimen in the Canadian National Collec-
tion was taken at Lloydminster, Alberta, well to the east of the Cordil-
lera, in the aspen parkland belt of the Prairie Provinces. These records
suggest that the species may occur in a series of disjunct populations
across central Canada.

In differing areas of the distribution, the species is in flight between
the middle of May and the middle of August. The period of adult ac-
tivity in any area is presumably governed by seasonal development; thus
at 1500 ft. in the Vernon area of southern British Columbia it flies during
the latter part of May, but at 6000 ft. in the Manning Park area it has
not been taken until early August.

Heliothis oregonica has been observed on several occasions ovipositing
in the heads of Castilleja spp. in both British Columbia and California.
The species is obviously not confined to Castilleja, however, because
partially grown larvae have been found feeding on the seed capsules of
Geranium sp.

2 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Behaviour

Heliothis oregonica is a day-flying species and has the compound eyes
greatly reduced as in other Heliothidinae with an exclusively diurnal
period of activity. On warm afternoons in the montane meadowlands
that constitute its usual habitat, the little moth flies swiftly and almost
invisibily from blossom to blossom. The female, with wings vibrating,
moves rapidly over a Castilleja head and pauses only momentarily, with
wings partially elevated, to deposit an egg.

On Castilleja the eggs are deposited singly on the bracts, on the lip of
the calyx, within the tube of the calyx or on the corolla. Three individ-
ually contined wild-caught females deposited a mean of 172 eggs, and
the maximum laid by a single female was 202. A few of these eggs
hatched on the fourth day after deposition but the majority hatched on
the fifth day. The newly hatched larva crawls down the tubular corolla
or bores through the calyx and corolla. Within the protective sheath of
the corolla, the young larva feeds on the stamens, pistil and developing
seed capsule. During the median stadia, the larva may usually be found
at the base of a bract, its anterior end buried in the adjacent blossom
on which it is feeding. Nearly mature larvae make much less effort to
conceal themselves, and may usually be found feeding from a partially
exposed position on the stalk. Both green and reddish-brown colour
phases of larvae are probably protected from predators by their re-
semblance to the green stems or to the red bracts and calyxes.

Forty-seven of 58 larvae, reared individually to the pupal stage, ma-
tured in six larval stadia, the remainder in five. At the cessation of feed-
ing, the larva tunnels into the ground to form a pupal cell. The species
overwinters in the pupal stage. Available data suggest that the species
is single-brooded throughout its range.

Descriptions of Stages

The following descriptions of immature stages were based on the
progeny of three females taken near Vernon, B. C., and on those of two
females taken near Tahoe Valley, Eldorado Co., Calif. The larvae were
reared individually at room temperature on the blossoms and seed cap-
sules of Castilleja spp. Rearing techniques employed were those out-
lined by Hardwick (1958). The estimate of variability following the
mean for various values is the standard deviation.

Adult (Figs. 1, 3). Head, thorax and abdomen with an inner coat of appressed

black scales and an outer coat of olive-grey hair-like vestiture; hair-like vestiture on
abdomen shorter and less dense than on head and thorax. Outer vestiture on under-

VoLUME 25, NuMBER 1

ee)

Figs. 1-6. Heliothis oregonica (Hy. Edwards) and its food plants. 1, 3, Adults,
Osoyoos, B. C., 3500 ft., and Vernon, B. C., 1200 ft.; 2, 4, dorsal and left lateral
aspects of ultimate-stadium larvae, Vernon, B. C.; 5, blossoms of Castilleja sp. on
which eggs have just been deposited; 6, pupae.

side of body paler than on upper side. Forewing olive-grey or olive-brown, variably
suffused with reddish-brown: Transverse anterior line sinuous, angling outward be-
tween costal and trailing margins; seldom well defined, often completely obliterated.
Median space cream, irrorated with olive or brown. Large, dark, usually globular
orbicular, claviform and reniform spots present in basal area of median space. Basal
space and basal area of median space variably suffused with olive or reddish-brown;
basal suffusion often extending outward to incorporate orbicular and claviform spots,
less commonly to also incorporate reniform spot. Transverse posterior line broadly
excurved around reniform spot, then essentially straight to inner margin. Subterminal
and terminal spaces concolorous, usually olive-brown, rarely suffused with reddish-
brown; the two spaces separated by a pale-yellow or pale-grey subterminal line.
Fringe concolorous with terminal space. Hind wing white marked with black; a
black basal area, a narrow black inner-marginal band, a broad black outer-marginal
band and a black blotch on disc. Black basal area usually extending outward to
incorporate at least part of discal blotch. Outer-marginal band with a white median

4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

patch. Fringe white. Underside of both wings pale grey marked with dark brown
or black, and with pale-grey fringes; outer area of each wing darker grey than
inner area. Forewing with prominent claviform, orbicular and reniform spots, and
a narrow subterminal band. Hind wing with a prominent discal blotch and a variably
expressed inner-marginal band; inner-marginal band forming an irregular patch at
anal angle and often produced to form at least a partial outer-submarginal band.

Expanse: 26.8 + 1.8 mm (48 specimens ).

Egg. Light green when deposited. Micropylar area becoming dull red on second
day after deposition. Reddish colouring becoming more intense on third day. Whole
egg becoming purplish-brown on fourth day after deposition; then turning dark
grey with larval head becoming visible through chorion a few hours before hatching.

Dimensions of egg: length, 0.566 + 0.061 mm; diameter, 0.577 + 0.023 mm (14
eggs).

First-Stadium Larva. Head and prothoracic shield black. Suranal shield dark
blackish-brown. Trunk dirty yellowish-grey in young larva, becoming clear pale
yellow in latter part of stadium. Setal bases and thoracic legs black.

Head width: 0.289 + 0.012 mm (25 larvae).

Duration of stadium: larvae maturing in five stadia, 4.4 + 0.8 days (11 larvae);
larvae maturing in six stadia, 4.2 + 0.6 days (47 larvae).

Second-Stadium Larva. Head, prothoracic and suranal shields usually dark
brown; head concolorous with, darker than or paler than prothoracic shield; head
occasionally rather light brown. Trunk dark dirty grey at beginning of stadium, be-
coming pale yellowish-green as larva increases in size; two pairs of pale dorsal lines
in many specimens. Setal bases and thoracic legs black.

Head width: 0.449 + 0.033 mm (25 larvae).

Duration of stadium: larvae maturing in five stadia, 6.9 + 1.0 days (11 larvae);
larvae maturing in six stadia, 6.4 + 1.5 days (47 larvae).

Third-Stadium Larva. Head fawn mottled with darker brown. Prothoracic shield
varying from green to pale fawn. Suranal shield poorly distinguished from trunk.
Trunk varying from dark greyish-green to pale bright green; two or three pairs of
rather prominent yellow lines on dorsal surface; midventral area greenish-grey, paler
than remainder of trunk. Thoracic legs fawn.

Head width: 0.745 + 0.069 mm (83 larvae).

Duration of stadium: larvae maturing in five stadia, 3.3 + 1.0 days (11 larvae);
larvae maturing in six stadia, 3.7 + 1.2 days (47 larvae).

Antepenultimate-Fourth-Stadium Larva. Head and prothoracic shield varying
from yellowish-fawn to greenish-fawn. Suranal shield essentially concolorous with
trunk. Trunk medium green, with a pair of dorsal and a pair of subdorsal, usually
prominent, yellow lines; three narrow wavy pale lines between dorsal and subdorsal
yellow lines. Lateral band prominent, the ventral half pale yellow or white, the
dorsal half pale green. Ventral region of trunk greyish-green, paler than dorsal
region. Thoracic legs varying from yellowish-fawn to greenish-fawn.

Head width: 1.12 + 0.07 mm (50 larvae).

Duration of stadium: 4.0 + 1.4 days (47 larvae).

Penultimate-Stadium Larva. Head varying from green to straw-coloured. Pro-
thoracic shield of similar colouring to head. Suranal shield poorly distinguished from
trunk. Trunk shades of green, in some specimens with a dull yellow cast. Mid-
dorsal band consisting of two or three narrow, irregular and broken lines. Subdorsal
area margined dorsally and laterally by yellow lines; median band of subdorsal area
consisting of two or three narrow, irregular and broken, green lines. Supraspiracular
area consisting of an irregular scrollwork of narrow green lines. Spiracular band white
or cream, stained with green dorsally. Ventral region greyish-green. Thoracic legs
translucent green or fawn.

VOLUME 25, NuMBER 1 5

Figs. 7, 8. Heliothis oregonica (Hy. Edwards), apical abdominal segments of
pupa. 7, Ventral; 8, right lateral.

A reddish-brown colour phase occasionally present among penultimate-stadium
larvae.

Head width: fourth-stadium larvae maturing in five stadia, 1.41 + 0.08 mm (23
larvae); fifth-stadium larvae maturing in six stadia, 1.74 + 0.10 mm (26 larvae).

Duration of stadium: fourth stadium of larvae maturing in five stadia, 5.8 + 1.4
days (11 larvae); fifth stadium of larvae maturing in six stadia, 4.6 + 1.3 days (47
larvae).

Ultimate-Stadium Larva (Figs. 2, 4). Larva occurring in several colour phases;
most larvae varying from light to medium green; a few larvae in shades of brown,
dull red, or even purple. Maculation in all these colour phases essentially the same.

Head and prothoracic shield green or yellow-fawn in green specimens; brown or
orange in brown or reddish specimens. Suranal shield essentially concolorous with
trunk. Middorsal band presenting a complex pattern formed by the remnants of three
or four narrow, badly broken, longitudinal lines. Subdorsal area similar to middorsal
band, flanked on either side by narrow light-yellow lines. Supraspiracular area similar
in appearance to middorsal band and subdorsal area. Spiracular band prominent;
cream, white or yellow; dorsal half of band suffused with ground colour of body.
Suprapodal area green or reddish-brown, mottled with grey. Midventral area vary-
ing from greenish-grey to brownish-grey.

Head width: 2.48 + 0.18 mm (25 specimens ).

Duration of feeding phase of ultimate stadium: larvae maturing in five stadia,
6.2 + 1.1 days (11 larvae); larvae maturing in six stadia, 6.6 + 1.6 days (47 larvae).

Duration of prepupal phase of ultimate stadium: 4.0 + 1.0 days (54 larvae).

Pupa (Figs. 6-8). Dark mahogany-brown with a green suffusion on thorax and
thoracic appendages. Spiracles on a level with general surface of cuticle; rims of
spiracles very weakly projecting. Abdominal segments 5, 6 and 7 extensively and
coarsely pitted; segments 4 and 8 less coarsely pitted and dorsum of segments 1, 2 and
3 rather finely pitted. Proboscis terminating between extreme apexes of wings. Cre-
master consisting of two long, moderately stout setae borne at the apex of thumb-
like prolongation of the tenth abdominal segment.

Length from anterior end to posterior margin of fourth abdominal segment: 9.0 +
0.8 mm (25 pupae).

Acknowledgments

I appreciate the assistance of my associate, Mr. E. W. Rockburne, who
measured the immature stages and drew the cremaster area of the pupa.

6 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Mr. John E. H. Martin of this Institute provided the photographs of
larvae and pupae.

Literature Cited

Epwarps, H. 1875. Pacific Coast Lepidoptera—No. 12. On some new species
of Noctuidae. Proc. Calif. Acad. Sci. 6: 132-139.

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

AMERICAN WHITE BUTTERFLIES (PIERIDAE)
AND ENGLISH FOOD-PLANTS

S. R. BowbEN
Redbourn, Herts, England

Six years ago (1964), when I offered notes on the rearing of sub-
species of Pieris napi L., I did so “in spite of their European bias.” What
I had in mind was the possible non-availability of recommended food-
plants in America, rather than their unsuitability for Nearctic subspecies.
However, even before the notes appeared, I had run into trouble with
Pieris virginiensis Edwards.

Most subspecies of the Pieris napi species-group will thrive on a va-
riety of cruciferous plants. Even bryoniae Ochsenheimer, which in its
Alpine habitats feeds almost exclusively on Biscutella laevigata L., shows
no obvious preference for this plant in captivity, and does quite well on
Alliaria petiolata Bieb. Other authors (e.g. Hovanitz & Chang 1962)
have attempted systematic experiments on the choice of food by Pieris
larvae (usually P. rapae L.), but it is difficult thus to demonstrate the
reality of host-species preferences, unless they are sufficiently marked to
over-ride possible differences in condition of the plants presented. The
experiments and observations now to be described were not systematic

in this sense, but by repetition and ruthlessness established certain con-
clusions.

Pieris virginiensis Edwards

May 1962.—Larvae hatching from Connecticut eggs of P. virginiensis
(kindly sent by Mr. S. A. Hessel) fed on Dentaria diphylla Michx. leaves
sent with them, but would do no more than nibble at English Alliaria,
Hesperis matronalis LL. and Cardamine pratensis L. All died. Later ex-

VoLtuME 25, NuMBER 1 7

perience suggests that a maintained supply of Cardamine might have
saved them.

May 1963.—Paired females from pupae received from S. A. H. laid ca.
200 eggs apparently indifferently on Dentaria diphylla and Alliaria; these
were divided into v(1), v(2) and v(3).

v(1)—F orty-three eggs on Alliaria were kept on this plant. Although the
larvae began eating the leaves, growth did not continue; many disap-
peared or died. After ca. 10 days only 2 survived; these were put on
D. diphylla and then on the very local English D. bulbifera L. One
nevertheless died; the other recovered and fed up normally (finally on
Cardamine ).

v(3)—One hundred and three eggs on Alliaria were further divided.
Twenty-four larvae, transferred to D. diphylla and thence to D. bulbifera,
suffered no check. All or most of the larvae remaining on Alliaria must
have fed slightly, judging by the condition of the leaves; after ca. 5-7
days the three very thin survivors were transferred to Dentaria. Two
nevertheless died, but one fed up well (finally on Cardamine ).

v(2)—Eggs on potted D. diphylla were transferred to D. bulbifera and
combined with the first 24 of v(3). Three larvae, just in the last instar,
were put on Hesperis. After some hours all 3 had left the leaves, which
they had not begun to eat. The following day the same 3 larvae were
again taken off Dentaria and were put on Alliaria for 24 hours; a very
little of this was eaten, but it was evidently unacceptable. The larvae
then fed up normally on Dentaria; the last 2 feeding (in Scotland) ac-
cepted Cardamine.

June 1964.—A further generation was bred from a single paired female.
Rearing losses were very high because (1) Dentaria had largely died
down, (2) the Cardamine-producing meadow had been sprayed with
weed-killer. Three of the butterflies emerged without diapause.

May 1965.—Broods were obtained from a pair from fresh S. A. H. pu-
pae (v) and by caging 2 such males with 3 females reared in 1964 (uw).

u—Egegs were all laid on Alliaria, none on Hesperis nor on Dentaria
bulbifera. Some larvae were fed almost entirely on D. bulbifera. Others
were transferred direct to Cardamine: after a slow start these grew well.
Still others were transferred first to Dentaria, thence to Nasturtium of-
ficinale R. Br. (watercress ), which was accepted.

v—Eggs were all laid on Alliaria, none on Hesperis. Initial rearing was
on Dentaria: one larva on Cardamine pupated ca. 7 days later than its

8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

contemporaries; two ca. 16 mm long were moved to watercress and of
these one at first refused but later accepted it. A few days later the re-
maining larvae were given watercress. Some larvae ate old Dentaria
petioles in preference to watercress; here, at least, the species of plant
over-rode its condition.

1966-67.—Six more broods were raised, largely on watercress. One in
1967 included a very few eggs laid on Hesperis but none on Cardamine.
One 1966 brood yielded a few “summer” specimens.

P. napi oleracea Harris

May 1964.—A pair, from New Hampshire pupae received from Mr.
Hessel, produced only 25 eggs. Alliaria was supplied as food, but several
larvae died and alternatives were tried: Sinapis arvensis L., Sisymbrium
officinale L. and Dentaria. No obvious preference for any of these was
shown, but there was no systematic experiment. The 9 butterflies ob-
tained after diapause were all small except one male.

July 1965.—Pairings from 1965 pupae received from S. A. H. produced
3 large broods. The young larvae were failing on Alliaria, so were trans-
ferred to Cardamine, and later to Sisymbrium and Hesperis. Two small
larvae were put on Alliaria only; very little was eaten and both larvae
were dead in 19 days. One larva which accepted Alliaria proved to be an
English waif.

P. napi marginalis Scudder

July 1966.—Stocks were raised from pupae (Saddle Mt. State Pk., Ore-
gon) received from Mr. C. W. Nelson. Watercress was used successfully
as larval food from the beginning. At first it was thought that Alliaria
(as well as the horseradish Armoracia) had been accepted by the first
batch of larvae, but some isolated larvae ate little or none of it. How-
ever, even larvae placed on Sisymbrium died and it became obvious that
part of this brood was unhealthy.

July 1967.—A later brood was given Cardamine + Alliaria. As the re-
maining Cardamine dried up several small larvae died, but no Alliaria
was eaten. Thereafter watercress and Sisymbrium were substituted.

Another brood all started to feed on Alliaria, but only one larva con-
tinued to do so, and this ultimately proved to be a Mediterranean stray.

June 1965.—A small batch apparently did well on Alliaria while
young, but were then given Sisymbrium (on which they made slow
progress ), Hesperis and Nasturtium. When about 10 days old the larvae
were divided: on Alliaria all 13 died; on Nasturtium 3 died and 13
pupated.

VoLUME 25, NuMBER 1 fe)

Hybrids
The breeding of hybrids of American subspecies will be described else-
where; the account is anticipated here only to state that American-British
hybrid larvae (both virginiensis 2 X napi 6 and napi 2 x oleracea é, as
well as the reciprocal hybirds of marginalis with Irish and British napi)
fed normally on British food-plants, including Alliaria, but virginensis x
oleracea hybrids rejected Alliaria.

Discussion

It has been known for a very long time (Verschaeffelt 1910) that
Pierinae feed almost exclusively on plants united in their possession of
mustard oils—Cruciferae, Capparidaceae, Resedaceae and Tropaeolaceae.
Verschaeffelt also successfully induced larvae to feed on almost pure
carbohydrates by flavouring them with juice containing natural mustard
oils. Such experiments have often been repeated since, with various de-
grees of sophistication, and the analyst can now easily distinguish the
various mustard-oil glucosides present in different species of Cruciferae,
eic.

The evolution of oligophagy has been discussed by Dethier (1954),
and the general subject of the co-evolution of butterflies and di-
cotyledonous plants has been admirably treated by Ehrlich and Raven
(1964). As in the quoted case of Papilionids on Aristolochiaceae, it would
appear that the Pierinae are a group which developed the ability to cope
with a class of toxic or repellent chemical products, which then became,
for them, attractants (among mammals, Hominidae appear now to be
attracted by many of these same toxins). While the Coliadinae and the
Dismorphiinae remained generally attached to Leguminosae, Pierinae
diversified in their chosen field and became increasingly selective in re-
spect of the combinations of volatiles demanded in their host-plants. We
may suppose that in a relatively stable environment specialization in any
one line normally increased until the population became confined to one
or two closely related species of plants, though the choice of these might
differ between conspecific populations in very different biotas.

If this is so, then we have in Pieris examples of “old” and “young”
species: P. virginiensis, P. ergane Geyer and perhaps P. bryoniae; P.
rapae L.. and European P. napi. Pieris ergane provides interesting paral-
lels with P. virginiensis. Another fragile butterfly of “relic” distribution,
it also has n = 26 chromosomes (Lorkovié 1941—see also 1968) although
belonging to the napi group which normally has n= 25; it also is re-

10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

stricted as to food, in this case to subspecies of Aethionema saxatile (L.)
R. Br. (Descimon 1966, Lorkovic 1968). Lorkovic’s females would lay
on none of 23 other crucifers.

When an oligophagous American “napi” is crossed with a polyphagous
European (in either direction), the hybrids are polyphagous. Whether
in these cases oligophagy is simply recessive was not determined. The
most restricted feeder, virginiensis, would have to be used to obtain un-
equivocal results, but after the F, only back-crosses to the supposedly
dominant napi were obtained.

The association between a butterfly species and its usual host-plant
requires that two conditions should be satisfied: (1) the female is at-
tracted to lay her eggs on or near the plant, and (2) the hatched larvae
eat it and go on eating it. As we have seen for P. virginiensis, females
may be super-normally attracted to a plant which the larvae refuse to
continue eating. On the other hand the wild females of P. bryoniae, by
laying their eggs almost exclusively on Biscutella (or locally on Arabis
halleri L.), conceal the readiness of their larvae to accept a wide range of
cruciferous plants.

If Alliaria petiolata were introduced to the American habitats of P.
virginiensis it is very probable that females would lay on it initially.
Subsequently either a larval variety would appear that tolerated Alliaria,
or the tendency to lay on Alliaria would be eliminated—the latter is
much the more likely sequel. In either case the cost of re-adaptation
might be very heavy.

It is clear that the inappropriate laying of captive females on Alliaria
is connected with the absence of this plant from the virginiensis habitat—
otherwise the undiscriminating behaviour could not have persisted.
Straatman (1962) has reported instances of four species of Australian
Lepidoptera laying in the wild on introduced plants; the general sequence
was that the larvae nibbled, refused to continue feeding and ultimately
died. In spite of the title of his notes, Straatman provides no evidence
that the plants were toxic to the larvae; it is more likely that his cases
were entirely analogous to our experiments with virginiensis.

The other “whites,” oleracea and marginalis, are not as restricted in
their choice of Cruciferae, though neither will continue to feed adequately
on Alliaria. P. n. oleracea has a wider range of known host-plants than
virgimiensis, and does include the latter's Dentaria diphylla. The sur-
vival of virginiensis where the two species are sympatric depends upon
its closer adaptation to life in the shady woods where Dentaria grows
(Rothke 1931, Voss & Wagner 1956). In England P. napi does lay on

VoLUME 25, NuMBER l ital

Dentaria bulbifera, but rarely, the habitat being rather too densely
wooded.

In the wild, P. virginiensis is reputed to be almost invariably univoltine,
though as we have seen it is potentially bivoltine. Since Dentaria di-
phylla is a “spring” plant, dying back early in June, butterflies which in-
herited a tendency to emerge in summer would have little chance of
reproduction if the larvae are in practice confined to Dertaria. A selec-
tive process will account for the failure of this rather southern species
to produce two generations in the year, which it could otherwise easily
do.

It would be of some interest to establish a colony of P. virginiensis in
Europe. If the attempt is made it should be in an area where Dentaria
is commoner than in England. It is to be expected that initially rather
free pairing with the local P. napi would occur, giving polyphagous hy-
brids. If Alliaria were present, part of the eggs would deposited on it,
and of these only the hybrids and back-crosses to napi would survive.
The colony might be wiped out before a barrier of sexual discrimination
could be built up. The experiment would be permissible, with such an
unaggressive insect as virginiensis.

Summary

Not all British Cruciferae on which European P. napi and bryoniae
subspecies can be raised are accepted by larvae of the American P. napi
oleracea, P. n. marginalis and P. virginiensis. Alliaria petiolata, on which
P. virginiensis actually lays preferentially, is initially attractive but larvae
will not continue to eat enough of it and die of inanition. However,
Cardamine pratensis and Nasturtium officinale are eaten, and the latter
is recommended. In addition, Hesperis and Sisymbrium can be used for
oleracea and marginalis.

European-American hybrids feed normally on Alliaria.

P. virginiensis is univoltine in the wild in consequence of its restriction
to a “spring” plant, Dentaria.

The choice of food-plants by Pierinae is discussed. In a relatively
stable environment, a species tends to become adapted to a very limited
range of hosts. For the larvae, though perhaps not for the laying female,
the optimum concentration of the attractants may be exceeded in certain
plants, so that feeding ceases; or alternatively a still intolerable repellent
compound may accompany the attractants.

The very specialised P. virginiensis has probably become increasingly
adapted, over hundreds of thousands of years, to an unchanging habitat;

1192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

now its archaic condition threatens its survival. On the other hand Pieris
napi has kept its evolutionary mobility and its subspecies have occupied
a variety of environments, some still changing rapidly; in that sense the
species has retained its youth.

Literature Cited

Bowpen, S. R. 1964. The maintenance for experimental purposes of form “sul-
phurea” of Pieris napi. J. Lepid. Soc. 18: 91-100.

Descimon, H. 1966. A propos de la plante nourricieére de Pieris ergane Geyer.
Alexanor 4: 207.

Deruier, V. G. 1954. Evolution of feeding preferences in phytophagous insects.
Evolution 8: 32—54.

Euruicu, P. R. anp P. H. Raven. 1964. Butterflies and plants: a study in co-
evolution. Evolution 18: 586-608.

Hovanitz, W. anp V. C. S. Cuanc. 1962. Three factors affecting larval choice
of food plant. J. Res. Lepid. 1: 51-61.

Lorxovic, Z. 1941. Die Chromosomenzahlen in der Spermatogenese der Tag-
falter. Chromosoma 2: 155-191.

1968. Systematisch-genetische und odkologische Besonderheiten yon Pieris
ergane. Mitt. schweiz. ent. Ges. 41: 233-244.

Roruke, M. 1931. Einige Notizen iiber Vorkommen und Lebensgewohnheit von
Pieris napi L. in Nordamerika. Int. ent. Z. 25: 262-263.

STRAATMAN, R. 1962. Notes on certain Lepidoptera ovipositing on plants which
are toxic to their larvae. J. Lepid. Soc. 16: 99-103.

VERSCHAEFFELT, E. 1910. The cause determining the selection of food in some
herbivorous insects. Proc. Acad. Sci. Amsterdam 13: 536-542.

Voss, E. G. anp W. H. Wacner. 1956. Notes on Pieris virginiensis .... hitherto
unreported from Michigan. Lepid. News 10: 18—24.

THE BRAZILIAN “CERCYONIS” (SATYRIDAE)

Ler D. MILLER
Allyn Museum of Entomology, 712 Sarasota Bank Building, Sarasota, Florida

and

THomas C, EMMEL
Department of Zoology, University of Florida, Gainesville

Much confusion has occurred in the delimitation of the genus Cercy-
onis Scudder (1875). Periodically various authors have sought to unite
these American butterflies with the Palearctic Satyrus Latreille (1810),
and Minois Hiibner (1819) whereas other authors have pointed out the
distinctness of Cercyonis. Miller (1968, pp. 99, 120) showed that the
Nearctic Cercyonis are members of the satyrine tribe Maniolini, and in
fact, the only American representatives of this basically Palearctic tribe,

VOLUME 25, NuMBER 1 13

whereas Satyrus and Minois structurally belong to the Satyrini. The only
representative of the Satyrini in the New World is Neominois Scudder
(1875), from the western United States. Emmel (1969) has described
the genus Cercyonis and by implication restricted it to the Nearctic.

The southern South American satyrid fauna has suffered from attempts
to relate the butterflies found there to Holarctic genera that were fa-
miliar to the northern hemisphere systematists that first described the
species. Many species belonging to the strictly American Pronophilini
and Euptychini were described and long retained in such northern genera
as Satyrus (Satyrini) and Epinephele (= Maniola: Maniolini). Such
insects are illustrated particularly in the Elina and Lymanopoda series
of the Pronophilini ( Miller, 1968, pp. 117-118). A similar situation exists
with regard to Cercyonis. Weymer (1912, pp. 228-230) listed the various
Nearctic Cercyonis and includes the South American glaucope (C. & R.
Felder) from southern Brazil and gustavi (Staudinger, 1897) from Bo-
livia. Forster (1964, p. 136) transferred the latter to the genus Argyr-
ophorus Blanchard (1852), a pronophiline. The more recently described
Cercyonis leuderwaldti Spitz (1931) must also be considered in this
review of the extra-Nearctic “Cercyonis.”

Should glaucope and leuderwaldti indeed be Cercyonis, a vast zoo-
geographic problem would arise: how did these butterflies get from
the Nearctic to the southern Neotropics without leaving intervening pop-
ulations, when did this occur and by what route(s)? Furthermore, to
which Nearctic species are these isolated populations related? The prob-
lem is purely academic, because leuderwaldti and glaucope are not even
in the same tribe, and neither is a member of the Maniolini, as is Cercy-
onis.

For comparison the venation, palpus, male and female forelegs and the
male genitalia of Cercyonis are given in Figs. 1-5.

Cercyeuptychia Miller and Emmel, new genus

Type-species: Cercyonis leuderwaldti Spitz. 1931. Rev. Ent. Sao Paulo 1: 46
(Brazil).

This genus is a member of the Euptychiini and conforms in the im-

portant characters with other members of the tribe, as defined by Miller
(1968, pp. 90-92). A formal description follows:

Eyes naked. Antennae short, between two-fifths and one-half length of wing;
club weakly developed, occupying distal quarter of antenna and slightly more than
twice thickness of shaft at its thickest point. Palpi (Fig. 7) about two and a half
times length of head, erect and slightly convergent at tips; third segment two-fifths
length of second, hairs of second segment less than twice greatest segmental width.

14 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-5. Cercyonis pegala (Fabricius). 1, C. p. alope (Fabricius), ¢ venation
(approx. 2X ), Connecticut, New Haven Co., Hamden (Allyn colln.); 2, same, palpus
(approx. 12x) (LDM slide M-2107); 3, same, ¢ foreleg (approx. 12x) (same
slide as Fig. 2); 4, C. p. texana (Edwards), 2 foreleg (approx. 12x) (LDM slide
M-2113); 5, C. p. alope, & genitalia (approx. 12x) (LDM slide M-2108).

Male foreleg (Fig. 8) reduced (forefemur-tibia-tarsus just over one-fifth length
of same segments of midleg) with monomerous, unspined tarsus; femur somewhat
longer than tibia. Female foreleg (Fig. 9) reduced, less than one-third length of
midleg, femur somewhat longer than tibia, with a pentamerous, clubbed tarsus bear-
ing spurs on the third and fourth subsegments. Ambulatory legs rather short, slen-
der; midtibia less than twice length of proximal midtarsal subsegment, slightly spiny
dorsad and with well-developed terminal spurs; midleg slightly shorter than hind
leg.

Forewing cell square-out, slightly excavate along mi-m., and about half length of
forewing costa. Forewing radial veins arising from cell in two branches, Rs and Mi
arising separately, M» arising slightly near M; than Ms, Cu: arising nearer Ms than
Cuz. Androconial patch of mealy and hairlike scales extending from 2A across cell
to Mi-M>, dentate distad in M:-M;. Sc and cubital stem inflated basally, 2A not.
(Fig. 6).

Hind wing cell straight, slight distal migration of ms-ms along Ms, and produced at
origin of Ms: length of cell to origin of M; about three-fifths length of wing to end
of Mz. Vein 3A slightly longer than Sc Ri, Ms and Cu, arising well separated and
Mz» arising nearer M; than Ms. (Fig. 6).

Male genitalia (Fig. 10) typical of those of many euptychiines (Forster, 1964),
especially as regards the free gnathos, a condition typical of many Euptychiini, but
not in Maniolini or Pronophilini. Genitalia similar to those displayed by genera

VOLUME 25, NuMBER l 15

Figs. 6-10. Cercyeuptychia leuderwaldti (Spitz). 6, ¢, venation (approx. 2 )
Brasil, D. F., Sobradinho, Brasilia (Allyn colln.); 7, palpus (approx. 12x) (LDM
slide M-2105); 8, ¢ foreleg (approx. 12x) (same slide as Fig. 7); 9, @ foreleg
(approx. 12x) (LDM slide M-1724); 10, 6 genitalia (approx. 12x) (LDM slide
M-2109).

Godartiana Forster (1964) and Praefaunula Forster (1964), but spined gnathos
typical of present genus.

The pattern of these butterflies (Figs. 16-19) is also reminiscent of
Godartiana and Praefaunula with the heavily striated under surface of
both wings and the lack of distinct bands that are prominent in most
other Euptychiini. The well-developed ocelli of Cercyeuptychia leuder-
waldti are only faintly indicated in any Godartiana, but well-developed
in some Praefaunula, and the wings of the present genus are rounded, as
in Praefaunula, not angular, as in Godartiana.

This remarkable genus is most closely related to Godartiana and Prae-
faunula, but quite distinct from both and immediately recognizable by
the spiny gnathos and much longer penis, and from Godartiana by the
wing shape. One of us (LDM) is working currently on the Euptychiini
and considers the present genus to be somewhat more advanced than
either Godartiana or Praefaunula, perhaps derived from one of them.
K. S. Brown (in litt.) states that the present genus has comparable habits
to Praefaunula armilla ( Butler ).

The name of the genus is feminine and derived from the similarity of
these butterflies to the Nearctic Cercyonis.

16 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 11-15. Pseudocercyonis glaucope (C. & R. Felder). 11, P. g. boenning-
hauseni (Foetterle), ¢ venation (approx. 2), Brasil, S. Paulo, Campos do Jordao
(Allyn colln.); 12, same, palpus (approx. 12x) (LDM slide M-2106); 13, same, 6
foreleg (approx. 16 ) (same slide as Fig. 12); 14, P. g. glaucope (C. and R. Felder),
2 foreleg (approx. 16%) (LDM< slide M-1725); 15, P. g. boenninghausemi, 6 geni-
talia (approx. 12 ) (LDM slide M-2110).

Pseudocercyonis Miller and Emmel, new genus

Type-species: Epinephele glaucope C. and R. Felder, 1867 [1864-1867]. Reise der
.. . Fregatte “Novara” ... Lep. Rhop., (3): 493-494; pl. 67, figs. 5, 6 (TL-
“Brasilia” ).

This genus is a member of the tribe Pronophilini and conforms in major
respects to the general characterization of that tribe by Miller (1968, pp.
110-114). A formal description follows:

Eyes naked. Antennae short, about two-fifths length of wing; club rather well
developed, occupying distal quarter of antenna, about three times thickness of shaft
and flattened at tip. Palpi (Fig. 12) about twice length of head, semi-porrect and
somewhat divergent; third segment less than one-fourth length of second, hairs of
second segment more than three times that of greatest segmental width.

Male foreleg (Fig. 13) greatly reduced (forefemur-tibia-tarsus only one-ninth
length of those segments of midleg), with a stubby, monomerous, unspined tarsus;
femur much longer than tibia. Female foreleg (Fig. 14) as aborted as that of male
with a monomerous, unspined tarsus bearing no spurs; femur much longer than
tibia. Ambulatory legs rather short and stubby; midtibia slightly more than twice

VOLUME 25, NUMBER 1 7,

length of proximal midtarsal subsegment, sparsely spiny dorsad and with well
developed terminal spurs; mid- and hind legs subequal.

Forewing cell square-cut, slightly excavate along mi-m2, and less than half length
of forewing costa. Forewing radial veins arising from cell in two branches, Rs and
M, approximate but separate at their origins, Ms arising midway between M; and
Ms and Cu: arising midway between M; and Cue. No distinct androconial patch.
Se greatly inflated at base, the cubital stem slightly inflated basad and 2A not at
allc@ris. 11).

Hind wing cell “stepped” distad with a definite distal migration of m2-m; along
M:, and produced at origin of Ms: length of cell to origin Ms slightly less than half
length of wing to end of Ms. Sc-R: subequal to 3A, Ms and Cu: arising well separate
and Mz arising somewhat nearer M; than M;, (Fig. 11).

Male genitalia (Fig. 15) quite distinct from those of either Cercyonis (Fig. 5) or
Cercyeuptychia (Fig. 10) but somewhat like those of Argyrophorus Blanchard, 1852
(Forster, 1964, pp. 135, figs. 168-169; Hayward, 1958, p. 254, fig. 43) and
Tetraphlebia C. and R. Felder, 1867 (Hayward, 1958, p. 257, figs. 45, 47), but the
longer, dorsally toothed penis immediately distinguishing the present butterflies.

The pattern of these butterflies (Figs. 20-21) is distinctive, but some-
what reminiscent of Tetraphlebia germainii C. and R. Felder, 1867 (Hay-
ward, 1958, pl. 4, fig. 161). Only by stretching the imagination could
glaucope be considered a Cercyonis, but it is not easily referred to Epi-
nephele (= Maniola) in which it was described, either.

This singular genus resembles at least the type-species of Tetraphlebia
superficially, but genitalically it is nearest Argyrophorus, the type of
which is the amazing aluminum colored A. argenteus Blanchard (1852)
from the mountains of Chile and Argentina. The venation of the present
genus is rather close to that of Argyrophorus (Miller, 1968, p. 112, fig.
279), differing in minor details; the present genus does not have a dis-
tinct androconial patch. The elongate, dorsally toothed penis of this genus
is characteristic. The present genus belongs to the Elina series of the
Pronophilini ( Miller, 1968, pp. 110, 117).

The generic name is feminine and refers to the fact that the type-
species was wrongly considered a member of the Nearctic Cercyonis by

many authors.

Discussion

The southern Neotropical “Cercyonis” are no more members of that
Nearctic genus than is true Cercyonis synonymous with the Palearctic
Minois. In fact, Cercyonis (Maniolini), Minois, (Satyrini), Cercyeupty-
chia (Euptychiini) and Pseudocercyonis (Pronophilini) are members of
totally different tribes within the Satyrinae. The similar facies and the
equivalent ecological niches shown by these four genera, as well as some
South African Dirini, suggest a possible adaptive advantage to a mor-
phological appearance such as shown by Cercyonis, etc., even though

18 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 16-19. Cercyeuptychia leuderwaldti (Spitz). 16, ¢ upper surface, Brasil,
Goias, Ponte Funda (Emmel colln.); 17, same specimen, under surface; 18, 9 upper
surface, Brasil, Goias, Orizona (Emmel colln.); 19, same specimen, under surface.

Figs. 20, 21. Pseudocercyonis glaucope boenninghauseni (Foetterle). 20, ¢, up-
per surface, Brasil, S. Paulo, Campos do Jordao (Allyn colln.); 21, same specimen,
under surface. Note: All figures approx. 1.5.

VoLuME 25, NuMBER 1 19

the butterflies are only remotely related. All appear to be scrub country
species, some in low country, as some Cercyonis and Minois, others at
high elevations, such as other Cercyonis and Pseudocercyonis. Such as-
semblages of ecological equivalents are not uncommon among the Sa-
tyridae, as demonstrated by the “Erebia series” of unrelated montane
butterflies, including the Holarctic Erebia Dalman (1816) (Erebiini),
the Lymanopoda series (Pronophilini) from the high Andes, Percno-
daimon Butler (1876) and other New Zealand Hypocystini and some
South African Dirini. Careful morphological examination is necessary
on members of supposedly cosmopolitan, and particularly pantropical,
groups to confirm or deny relationships that have all too long been
taken for granted.

Acknowledgment

We should like to thank Dr. Keith S. Brown, Jr., of Rio de Janeiro,
Brazil, for providing material and observations on the ecology of the
south Brazilian “Cercyonis.”

Bibliography

Emme, T. C. 1969. Taxonomy, distribution and biology of the genus Cercyonis
(Satyridae). I. Characteristics of the genus. Jour. Lepid. Soc. 23: 165-175.

Forster, W. 1964. Beitrige zur Kenntnis der Insectenfauna Boliviens XIX.
Lepidoptera III, Satyridae. Veroff. Zool. Staatssamml. Miinchen 8: 51-188.

Haywarp, K. J. 1958. Satiridos argentinos (Lep. Rhop. Satyridae) III. Guia
para su clasificacion. Acta. Zool. Lilloana 15: 199-295.

Mmuuier, L. D. 1968. The higher classification, phylogeny and zoogeography of
the Satyridae (Lepidoptera). Mem. American Ent. Soc. 24.

WeyYMER, G. 1910-1912. Satyridae. in Seitz, A. Die Grossschmetterlinge der
Erde, vol. 5 (Die Amerikanische Tagfalter). Stuttgart.

CONSUL PANARISTE (NYMPHALIDAE) IN VENEZUELA

I secured two fresh males of Consul panariste (Hewitson) on 5 and 6 February
1968 while collecting in the Venezuelan Andes with Albert Gadou of Caracas. They
were taken on banana bait at approximately 1000 meters elevation on the Barinitas
to Santa Domingo road in the state of Barinas. This is a humid tropical forest situa-
tion, transitional to cloud forest. Albert reported having taken previous examples of
the species in the same location.

Comstock (1961. Butterflies of the American Tropics: The genus Anaea, p. 188)
stated that Consul panariste is known to occur only in Colombia. C. panariste has
been traditionally placed in the genus Anaea, before Comstock allied it with Consul
fabius (Cramer) (= Protogonius hippona Fabricius). Although Comstock con-
sidered Consul to be a subgenus of Anaea, contemporary usage usually elevates the
subgenera in his monograph to generic rank.

Jonn H. Masters, Lemon Street North, North Hudson, Wisconsin.

20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

SYMBIOTIC RELATIONSHIP OF AN ECUADORIAN SKIPPER
(HESPERIIDAE) AND MAXILLARIA ORCHIDS

THomas C. EMMEL
Department of Zoology, University of Florida, Gainesville

On the eastern side of the Andes in Ecuador lies the small town of
Puyo, the terminal point for biologists wishing to travel into the Oriente
or Amazonian Basin. Lush rain forest and rich butterfly faunas are found
both in the Andean foothills to the west and the lowlands to the east.
The town itself is at an elevation of 3,650’. On an expedition to this re-
gion in August and September 1969, I observed a most interesting pol-
linating relationship, apparently mutualistic, between a large skipper but-
terfly species, Perichares philetes dolores (Reakirt), and a native arboreal
orchid Mazxillaria ontoglossom.

At the orchid gardens of Mr. Joseph Brenner, manager of the Hostoria
Turingia in Puyo, a number of these Mavillaria ontoglossom orchids from
the surrounding forest are under cultivation. On September 8, I observed
a female P. philetes dolores visiting the flowers of these plants around
10 a.m. Mr. Brenner noted that this species of skipper, quite distinctive
in size and coloration (see Fig. 1), visits these flowers daily (observa-
tions of about 18 montlis).

This species of skipper seems adapted specifically for feeding from
and pollinating flowers like those of this long-tubed species of Mazxillaria
orchid. It has an extraordinarily long proboscis (Fig. 2). In the female I
examined (forewing length of 27.0 mm, and a body length from head to
tip of abdomen of 22.0 mm), the tongue was fully 43.0 mm long when
extended. This long proboscis can be flexed at any or several points by
the insect, and seems perfectly adapted for probing through the exten-
sively convoluted corolla tube to reach the nectaries.

The orchid deposits several pollinia (sticky packets of pollen grain)
on the proximal end of the proboscis and these are carried on to the next
flower the butterfly visits. Examination of the orchid blossoms showed
that the extremely long, convoluted, and constricted corolla throat pre-
vents entrance by bees or flies and that the nectar can only be reached
by a very long-tongued insect. This skipper species was never observed
visiting other orchids or flowers of any kind.

Thus it is suggested that this is a mutualistic symbiosis, with the
skipper species having a guaranteed nectar source barred from other
insects, and the orchid being pollinated by a specific pollinator which

VoLuME 25, NuMBER | 21

Fig. 1. Perichares philetes dolores (Reakirt), dorsal and ventral surfaces of
female.

apparently visits, at least primarily, these flowers of Mavillaria ontoglos-
som in the Puyo area of Ecuador. Since the skipper does occur north
through the Neotropics (Seitz 1913), it undoubtedly uses a variety of
orchids or other plant species in the various parts of its range, and it

Fig. 2. Perichares philetes dolores perched on lip of the orchid Maxillaria onto-
glossom, preparatory to feeding. The proboscis is normally extended straight down-
ward at about a 30° angle from the body plane during feeding (distal portion re-
curved in above sketch to conserve space).

92, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

would be interesting to have detailed data on specificity (or lack of it)
in these other areas.

Acknowledgments

I thank Dr. Lee D. Miller, Curator of the Allyn Museum of Entomol-
ogy, Sarasota, Florida, for identification of the skipper species, and the
Organization for Tropical Studies for their support of my 1969 research
expedition to Ecuador.

Literature Cited

Seitz, A. (editor). 1913. The Macrolepidoptera of the World. Vol. V. The
American Rhopalocera. Stuttgart.

FIELD TECHNIQUES FOR INVESTIGATIONS OF POPULATION
STRUCTURE IN A “UBIQUITOUS” BUTTERFLY

PETER F. BRUSSARD
Langmuir Laboratory, Cornell University, Ithaca, New York

Because information about population structure is necessary for a
proper understanding of the ecology, evolution, and geographic varia-
tion of any species of Lepidoptera, it is surprising that there have been so
few effective population studies of these organisms. The investigations
of Dowdeswell, Fisher, and Ford (1940, 1949), Turner (1963), Keller,
Mattoni, and Seiger (1966) and Ehrlich (1965), among others, are notable
exceptions. These studies have all documented essentially similar popula-
tion structures, and have led to the belief that the subdivision of butterfly
species into small isolated or semi-isolated populations with limited inter-
change of individuals is a general rule.

Studies conducted in 1962, and 1967-1969 on the satyrine Erebia
epipsodea Butler, however, have revealed a population structure quite
different from those previously reported (Brussard & Ehrlich 1970a,
1970b). We have now determined that the population studied in the vi-
cinity of Rocky Mountain Biological Laboratory (RMBL), Gunnison
County, Colorado may cover hundreds of square kilometers. There is a
great deal of individual movement, and, although these butterflies are
capable of recognizing and leaving ecologically unsuitable areas, these
areas are not barriers that subdivide the population into smaller units.

VoLUME 25, NuMBER 1 93

Since the population structure shown by E. epipsodea may be more
typical of Lepidoptera in general than the species previously studied,
the purpose of this report is to describe in detail the field techniques used
in determining this structure. Details of data analysis are reported else-
where ( Brussard and Ehrlich, 1970a).

When the study began, it was expected that each subalpine meadow
or sagebrush flat supported a separate and distinct population, rarely ex-
changing individuals with adjacent areas, and that the strips of aspen
and alpine fir that divided the meadows into discrete units would also
serve as the boundaries of the butterfly populations. A capture-mark-
recapture program initiated in 1962 used several meadows in the vicinity
of RMBL as individual sampling areas. Analysis of the results of this
program revealed that out of 735 butterflies collected, marked, and re-
leased in 5 localities, less than twenty percent were ever recaptured—
far less than one would expect from examining the results of similar
studies. Nevertheless, approximately eighty percent of the butterflies
recaptured were taken in the area of previous capture. There was no
way to determine whether these paradoxical results were caused by high
mortality, large population size, or undetected migration.

Additional complications were caused by the short flight season of
E. epipsodea in this area (approximately three weeks, the exact dates
varying somewhat with elevation) and the unpredictable weather charac-
teristic of early summer in the high, mountainous regions of Colorado.
In late June and early July the mornings are often cool and cloudy, fol-
lowed by a brief period of relatively clear sky from mid morning to
early afternoon. Clouds and thundershowers usually follow from mid
afternoon to early evening. Since it had been shown that E. epipsodea
does not fly at all in temperatures below 14° C and in any numbers below
18-19° C, nor does it fly when a strong wind is blowing or when the sun
is obscured by clouds (Brussard & Ehrlich, 1970c), the time available
for collecting in any one day was limited. Any capture-recapture analysis
is based on the assumption that marked animals have had time to mix
freely with the remainder of the population before recapture. Thus, in
order to study the population structure of this insect, it was decided that
large numbers of E. epipsodea must be marked, released, and recaptured
in a systematic manner within the limited period of time available.
Furthermore, the sampling program must be designed to (1) give ac-
curate population size estimates and (2) provide quantifiable informa-
tion on inter-area movement. The development of a systematic sam-
pling program in 1967 gave results which suggested the true nature of the

94 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

population structure of this insect. Modifications of this program in 1968
verified and refined the results of the previous year.

Because of the restrictions imposed by the alpine environment, and
the large number of butterflies flying in the study areas, it was necessary
to develop techniques for rapid handling of the insects in the field. A
rapid marking technique was needed, since methods previously described
(see, for example, Dowdeswell, Fisher, and Ford 1940 and Ehrlich and
Davidson 1960), are far too slow or involve too many people in the
marking process. It was found that the “Sharpie” marking pen, manu-
factured by Sanford’s, was the most satisfactory marking device. Be-
cause of the fine point, good ink flow and rapid drying, one person using
this pen could mark the butterflies quickly and efficiently. Furthermore,
the “Sharpie” is available in several colors, each of which can be dis-
tinguished from the other after application. Although the color black
tended to dry on the tip of the instrument if it were not instantly capped
after applying a mark, this difficulty was not experienced with any of the
other colors.

Weather permitting, collecting began around 0900 and continued for
approximately one hour. The butterflies were netted and placed in in-
dividual glassine envelopes which were stored in styrofoam ice chests
kept at convenient spots within the study areas. After all butterflies ob-
served in an area had been captured, they were removed from the en-
velopes with forceps and examined. The recapture event was recorded
for marked individuals; freshly caught ones were assigned a number and
marked. In addition to date and area of capture, sex and condition were
recorded at the time of initial capture and at every subsequent recapture.
Damaged individuals or those exhibiting abnormal flight behavior upon
release were not returned to the population. There is no evidence that
there were any differences in behavior or probability of recapture be-
tween marked and unmarked individuals.

The numbering technique used was a modification of the method de-
scribed by Ehrlich and Davidson (1960). The modifications included
adding additional spots near the base of each wing which were assigned
values of 100, 200, 400 and 700, increasing the number of individuals
that could be marked, per color, to 1000 (Fig. 1). Because of this, and
since the “Sharpie” is available in several colors, it was possible to give
individual numbers to all butterflies marked in 1968. (In 1967 most.
insects were individually numbered; however, some were marked with
a code pattern indicating area and date of capture.) The advantages of
individually numbering all butterflies handled during a flight season are

VOLUME 25, NuMBER lI

bo
Ol

oW/,

Ze 810

Fig. 1. Modification of Ehrlich and Davidson’s (1960) marking system; 1000
butterflies per color can be marked in this fashion.

obvious; considerably more information is accumulated on individual
movement, and many more partial histories of recaptured insects can be
compiled.

The manner of release of animals as vagile as butterflies assumes great
importance in population studies. In order to evaluate the possibility
that an abnormally high level of activity occurred subsequent to re-
lease, causing excessive dispersal out of the study areas, on two occa-
sions the butterflies were held and released in late afternoon under
cloudy conditions. Collecting was delayed the following morning giving
the butterflies an opportunity to mix freely with the remainder of the
population prior to recapture. The percentage of recapture for that
day was then compared to both the day preceding and the day following
when normal release procedures were followed. Since there was no sig-
nificant difference (p >> .05), it was assumed that the marking pro-

26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Summary of Capture-Recapture Results, 1967

Number of % of Total
Days Marked & individuals individuals number of
Locality used. Sex released recaptured recaptured recaptures*
M 204 14 6.4 14
Visquene Meadow 7
F 13 0 0) 0
M~ 354. 123 34.7 180
Cemetery ILL
F Bn) 4 1255
Bench Meadow M 147 4.1
Area 4
F 3 0 0 0
Naked Hills M JEL 19 17 19
Peninsula 4.
F 6 0 0
M 66 Z, 3.0
Wilson Ranch 4
Ie 3 0 0 0
M Pali IPALL 57.3 142
Gothic (combined ih
data for two F 2, D2, 31.9 Aas)

adjacent plots )

* includes multiple recaptures

cedure did not cause excessive dispersal. An attempt was also made to
release the butterflies at points scattered throughout the study area to
facilitate free mixing with the rest of the population.

When the capture-recapture program was resumed in 1967, six locali-
ties were chosen on the basis of accessibility and apparent abundance of
butterflies. In these localities 1222 butterflies were marked and released
and an additional 1259 butterflies were captured and examined in pe-
ripheral areas. Examination of these data (Table 1) showed that (1)
large plots surrounded by even larger expanses of ecologically suitable
habitat gave exceedingly small recapture percentages, no matter how
long they were sampled (Visquene Meadow, Bench Meadow, Wilson
Ranch). (2) Large plots isolated or semi-isolated from other suitable
areas gave higher recapture percentages, and the percentages improved
with the length of the sampling period. (Cemetery, Naked Hills Penin-
sula). (3) Two small, adjacent plots, semi-isolated from other areas of
suitable habitat and intensively sampled for relatively long periods of
time (Gothic) gave satisfyingly high recapture percentages; furthermore,
the positioning and size of these plots made it possible to calculate move-

VOLUME 25, NuMBER 1 27

ment of individual butterflies between them. However, when these
data were further analyzed in order to get daily population size estimates,
it was found that the estimates were highly erratic and not in keeping
with field observations. In contrast to the erratic fluctuations in popula-
tion size estimations from data from the small plots, those estimates made
for the larger area (Cemetery), were considerably smoother and tallied
more with field observations. The analytical method employed for these
estimations was the stochastic model developed by Jolly (1965). The
possibility that the difficulties in estimation, especially in the smaller
plots, might be reduced by employing a different method of analysis
was considered, although the advantages of stochastic models in general
and Jolly’s method in particular have been pointed out by Southwood
(1966). Since the possibility existed that the Jolly method might not be
entirely suitable for E. epipsodea, these results were compared with re-
sults derived from the deterministic models of Dowdeswell, Fisher, and
Ford (1940) and Bailey (1951, 1952). In each case the estimates from
these deterministic models were consistently 40% to 60% lower than the
Jolly estimations. Indeed, on several days, the daily population size esti-
mates based on Bailey’s method were lower than that day’s catch.

It was concluded, therefore, that the Jolly method was the most suit-
able analytical tool and that the size of the study plot assumed consider-
able significance in the estimation of population parameters. The 1968
sampling program, therefore, was designed to gather additional data
amenable to analysis by this method.

It had been established during the 1967 season that one worker could
usually capture all the E. epipsodea flying in an area of approximately 1
hectare (2.5 acres) in one hour. During peak flight season this was ap-
proximately 75 insects (Brussard and Ehrlich, 1970a). It was also found
that one collector could not adequately cover areas too much larger
than 1 ha during the time available for collecting and capture enough
butterflies to ensure that recapture percentages were high enough for
analysis. In order to utilize an area large enough to give reasonable
population size estimates, we employed, in 1968, three field workers for
the Cemetery area, trimming the area somewhat (from 7.6 ha to 5.5 ha)
to ensure thorough coverage. Since direct evidence of movement of but-
terflies from site to site along with evidence of re-immigration would
provide the main support for the type of population structure that has
been advanced for E. epipsodea (Brussard and Ehrlich, 1970a), it was
also necessary to establish smaller study plots that could be simultane-
ously covered by each field worker so that these types of data could be

28 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

accumulated. To this end, the Gothic area was expanded to include 3
contiguous plots of approximately 1 ha which were sampled simultane-
ously for 11 days. It was then possible to document thoroughly move-
ment between areas and to estimate its magnitude. The 1968 results
confirmed that dispersal and re-immigration did explain the difficulties
encountered in obtaining the population size estimates in the small plots
(Brussard and Ehrlich, 1970a ).

The techniques developed here appear to be very useful in studies
designed to establish density and population structures of diurnal Lepi-
doptera, especially those which appear to be more or less continuously
distributed in their habitat, without ecological “barriers.” The results of
extensive studies such as these are a prerequisite for (and may largely
determine the methods to be used in) intensive studies designed to
delineate those factors that cause or regulate fluctuations in population
size. Many more data pertinent to these phenomena are needed before
the important controversy concerning density dependent or density in-
dependent population size regulation can be resolved.

Acknowledgments

This research was supported by funds from National Science Founda-
tion Grant GB-5385 and 8038. The use of the facilities of the Rocky
Mountain Biological Laboratory is gratefully acknowledged. Paul R.
Ehrlich and Robert H. Whittaker read the manuscript and made many
helpful suggestions; Jerry and Susan Downhower made the illustration.

Literature Cited

Baitey, N. T. 1951. On estimating the size of mobile populations from recapture
data. Biometrika 38: 293-306.

1952. Improvements in the interpretation of recapture data. J. Anim.
laeell, Zils IOO—1D7/.

Brussarp, P. F. anp P. R. Enric. 1970a. The population structure of Erebia
epipsodea (Lepidoptera: Satyrinae). Ecology 51: 119-129.

1970b. Contrasting population biology of two butterfly species. Nature

227: 91-92.

1970c. Adult behaviour and population structure in Erebia epipsodea
(Lepidoptera: Satyrinae). Ecology 51: 880-885.

DowvesweL1, W. H., R. A. Fisner, AnD E. B. Forp. 1940. The quantitative study
- as in the Lepidoptera. 1. Polyommatus icarus Rott. Ann. Eugenics

——— 1949. The quantitative study of populations in the Lepidoptera. 2. Maniola
jurtina L.. Heredity 3: 67-84. |

Kuruicw, P, R. 1965. The population biology of the butterfly Euphydryas editha.
II. The structure of the Jasper Ridge colony. Evolution 19: 327-336.

FuruicH, P. R., ann S. E. Davipson. 1960. Techniques for capture-recapture
studies of Lepidoptera populations. J. Lepid. Soc. 14: 227-29.

VOLUME 25, NuMBER 1 29

Jotty, G. M. 1965. Explicit estimates from capture-recapture data with both
death and immigration-stochastic model. Biometrika 52: 225—47.

KELLER, E. L. Jr., R. H. T. Matroni, AND M. S. B. Sricer. 1966. Preferential
return of artificially displaced butterflies. Anim. Behav. 14: 197-200.

SoutHwoop, T. R. E. 1966. Ecological Methods. Metheun & Co., London.

Turner, J. R.G. 1963. <A quantitative study of a Welsh colony of the large heath
butterfly, Coenonympha tullia Muller (Lepidoptera). Proc. R. Ent. Soc. Lond.
38: 101-112.

LEPIDOPTERA IN THE UNPUBLISHED FIELD NOTES OF
HOWARD GEORGE LACEY, NATURALIST (1856-1929)!

Roy O.? anp C. A. KENDALL
135 Vaughan Place, San Antonio, Texas

Howard George Lacey was born 15 April 1856 at Wareham, Dorset,
England. His elementary education was received at Charterhouse,
Hampshire; he studied at Frankfurt, Germany and later received a B.A.
degree from Cains College, Cambridge. Although educated for the min-
istry, he soon gave up this career for that of naturalist. At the age of 26
he came to the United States, arriving in New York 17 June 1882, and
proceeded to Kerrville, Texas where he arrived 30 June. Here he bought
a ranch in the hill country about seven miles southwest of Kerrville on
Turtle Creek (Figure 1) where he spent the next thirty-seven years rais-
ing livestock and studying the natural history of the area.

Lacey was a member of the Bournemouth Scientific Society, San An-
tonio Scientific Society, National Geographic Society, American Audubon
Union, and American Ornithologists’ Union. He collaborated with the
Smithsonian Institution, U. S. Department of Agriculture, and other in-
stitutions for the advancement of natural science.

So far as can be determined, he published no scientific papers. In-
stead, he chose to collect biological specimens which were sent to various
contemporary taxonomists and natural history museums. Many of his
field collections and observations were recorded in a ledger which is the
basis for this account. His ledger is now in the Witte Memorial Museum
Library, San Antonio, Texas placed there by his sister Miss Beatrice Lacey.

1 Contribution No. 198, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville.

2 Research Associate, Florida State Collection of Arthropods, Division of Plant In-
dustry, Florida Department of Agriculture and Consumer Services.

30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Pda SO eh

IGP por 2 48
function Su 6
igs

Mig. 1. A, Kerr and surrounding Texas counties; B, location of Howard G.
Lacey's ranch in relation to Kerrville, Texas.

VOLUME 25, NUMBER 1 31

Some of the botanical and biological specimens named in his honor
are: Quercus laceyi Small, Strymon laceyi Barnes & McDunnough, Grap-
tolitha laceyi Barnes & McDunnough, Logoa laceyi Barnes & McDun-
nough, Peromyscus boylei laceyi Attwater, and Reithrodontomys laceyi
Attwater. Only the last two were mentioned in his field notes.

Laceys primary interests were birds and mammals of which he made
numerous records. Later he became interested in Lepidoptera. The first
and last recorded notes pertaining to Lepidoptera were 28 October 1882
and 23 April 1919 respectively. On 10 April 1898 he wrote: “Saw indigo
bird and caught a few butterflies for Dr. Barnes. I intend to study the
butterflies of the neighborhood from this date.”

His collecting was not limited to butterflies. Many moths were col-
lected, apparently at lantern light. Although most of the specimens were
sent to taxonomists and institutions, he also accumulated a personal col-
lection. On 16 May 1902 he wrote: “W. G. Dodge came over from Kerr-
ville to see the butterflies.”

Preparatory to moving back to England (8 Aug. 1919), Lacey left his
local Lepidoptera collection with C. Stanley Coppock, an intimate friend,
who had also come from England and settled at Kerrville a few years after
Lacey. In 1925, Mrs. Ellen S. Quillin, then director of Witte Memorial
Museum, attempted to obtain the collection for the Museum. Still later,
his field notes were acquired. Excerpts from correspondence between
Mrs. Quillin and Miss Beatrice Lacey, sister of Howard, disclose what

happened.
On 25 March 1934 Miss Lacey wrote: “I have been thinking things over
since my last letter to Mr. Coppock. .. . I feel Howard would like me to

give his notes and diary to the Witte Memorial Museum if you say they
will be of use, and will be taken care of. It hurts to part with them, but I
would rather feel they are of use when I have passed over and Howard
would like them to be of use I feel sure. I ask you to accept them in his
name and mine with all good wishes.”

Mrs. Quillin’s reply of 11 April 1934 stated in part: “I received your
letter this morning in which you are transferring to us as a gift the life-
time notes of your brother. I certainly believe you are doing the right
thing and that Mr. Lacey would approve were he here, because when
Mr. Lacey visited this Museum, at that time only two school rooms
thrown together to make a Museum, the last time he was in Texas [1925]
and the Attwater collection had just been added, he expressed his regret
that the butterfly and moth collection he had made could not go with the
Attwater collection. It seems that when he left his collection with one

32, JouRNAL OF THE LEPIDOPTERISTS SOCIETY

of the Coppock children [1919] bugs got into some of it and other
specimens were sold off. He said that the selling was with his consent
inasmuch as he had then given up Texas ever getting a Museum in which
to place them. That summer [1925] after he left I tried to locate the
collection and found that it was held for sale. Mr. Lacey had given me
only his verbal wish and I, therefore, could not push the matter.

“We have had made a typewritten copy [300 pages] of Mr. Lacey’s
notes which I bound last week so that scientists of note might use them
for reference without in any way fingering or tearing the originals. The
original books will be taken to be bound this afternoon and from then on
will be placed under lock and key in my own office.”

Lacey's field notes are a veritable data bank on the local flora and
fauna. Most of his collecting was done at or near his ranch. Each time
he ventured far from the ranch an entry was made in his field notes. For
those museums having specimens collected by Lacey with dates, but
without locations, the following may be of help. Dates are inclusive.

28 February—27 October 1888, trip to England and return.

8 May 1893-10 April 1894, trip to England and return.

3-23 November 1896, hunting trip on Nueces River and Frio Divide.
[Apparently Uvalde County Texas. |

15-26 July 1898, fishing trip to Paint Creek. [This creek empties into
the South Llano River. Most of it lies in northeast Edwards County
Texas. |

25-26 June 1899, to San Antonio [Bexar County Texas].

27-[28] June 1899, went to the Medina [River] and back the next day.
[Apparently in Bandera County Texas. |

26 July—5 August 1899, went to the Llano [River] fishing and had a
very jolly trip. I got no new butterflies but took several very dark speci-
mens of T. damon [Callophrys gryneus castalis.| [Apparently Kimble
County Texas].

15 November-14 December 1899, our usual hunt on the Frio [River].
[ Uvalde County Texas. ]

7-19 July 1900, fishing on Llano River. As there were no flowers out
[ got only a few [butter] flies. On July 17th when we started home the
South Llano was up about five feet. [Probably Kimble County Texas. |

25-30 July 1901, got back to town after a short trip to the Llano [River]
with Carr and company. We camped a few miles above London. Elada
|Texola elada| was about the only butterfly about. [London is in north-
east Kimble County Texas. ]

VOLUME 25, NuMBER Il 33

27 August 1901-29 March 1902, trip to England and return. Got back
to the ranch little before sundown.

13-27 July 1902, fishing trip to Llano [River] camped near head of
South Llano River. [Edwards County Texas. |

10-31 July 1904, fishing on Llano River. Few [butter] flies as bloom
was very scarce. [Probably Kimble County Texas.]

17 July—2 August 1907 (19-31 July in camp) on fishing trip to the San
Diego River in Mexico via San Antonio and Del Rio, Texas.

15-31 July 1909, fishing trip to Devils River. [Val Verde County
Texas. |

23 November-6 December 1916, hunting trip to Paint Creek [Ed-
wards County Texas].

16 July 1919, sold my ranch and stock. After selling the ranch, I put in
a day or two loafing around Kerrville and then spent a week on Paint
Creek swimming near the spring. Eagles, I think, used the nest on the
bluff above the springs and the herons seem to have increased in num-
bers; nine nests in the big tree. [Apparently this was one of Lacey’s fa-
vorite spots. ]

8 August 1919 he left Kerrville, Kerr County, Texas for his native home
in England. Still fascinated by the wonderful Hill Country of the Ed-
wards Plateau and drawn by the devoted fellowship of old friends, he
returned to Kerrville in summer of 1925 for a short but last visit, health
failing. He died 5 March 1929 at the home of his sister, Miss Beatrice
Lacey, 50 Wellington Road, Bournemouth, England.

In the following list of Lepidoptera mentioned in Lacey’s field notes,
arrangement generally follows dos Passos (1964) for the skippers and
butterflies, and McDunnough (1938) for the moths. The italicized name
is the species most likely collected or observed by Lacey; often he re-
corded only the species name. The exact citation given by him is shown
below the complete scientific name. Following this are the dates and
locations of specimens collected (c) or observed (s). If he gave a quan-
tity, this too is shown. Example: (c-3) or (s-1) means he collected three,
or saw one. Other pertinent remarks from his field notes are also given.
Statements in brackets are those of the authors. If a specific county is
not given, it may be assumed that Lacey collected or observed the speci-
mens at or near his ranch located about seven miles southwest of Kerr-
ville, Kerr County, Texas.

Of the 144 species of Lepidoptera recorded by Lacey, several are no
longer to be found in the Kerrville area, probably due to extensive grazing
by domestic sheep and goats. A few species could be considered only

34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

strays even at the time collected. He recorded migratory flights of Krico-
gonia lyside, Libytheana bachmanii, and Danaus plexippus.

Lepidopterous species collected by Lacey and included in the list but
not recorded in his field notes are: Tmolus azia (Hewitson), Strymon
alea (Godman & Salvin), Chlosyne endeis (Godman & Salvin), Cale-
phelis rawsoni McAlpine, Graptolitha laceyi Barnes & McDunnough, and
Lagoa laceyi Barnes & Dunnough.

Megathymidae

Megathymus coloradensis kendalli H. A. Freeman 1965. “M. yuccae’. 8-IV-
1900 (c), 15-IV-1900 (c), 15-III-1901 (s), I7-III-1901 (c-14 ), 18-III-1901 (s-2),
19-III-1901 (c), 20-III-1901 (c), 22-III-1901 (s), 12-IV-1901 (s), 18-IV-1901 (c),
4-IV-1902 (s), 7-IV-1902 (c), 22-IV-1902 (c), 30-IV-1902 (s), 1-IV-1903 (s),
26-IV-1903 (c-19), 10-V-1903 (s), 24-III-1910 (s), 2-V-1913 (c), 4-V-1913 (c).
[Have recent records, but scarce. ]

Hesperiidae

Panoquina ocola (Edwards) 1863. “Ocola”’. 19-VIII-1907 (c-3), have not seen
it since 1899. [Should be found occasionally. ]

Lerodea eufala (Edwards) 1869. “Eufala’”. 18-III-1900 (c), didn’t see eufala
all last year, the year before it was common; 11-IV-1901 (c). [Have recent rec-
ords. ]

Amblyscirtes aenus aenus Edwards [1878]. “Amblyscirtes aenus’. 23-IV-1899
(c-3), 23-IV-1903 (c), 26-IV-1903 (c). [Have recent records. ]

Amblyscirtes oslari (Skinner) 1899. “Oslari”. 15-IV-1900 (c). [Have recent
records. |

Amblyscirtes nysa Edwards 1877. “Nysa”. 24-III-1900 (s), 15-IV-1900 (c),
1-IX-1900 (c), 4-IV-1901 (c), 18-VI-1902 (ec), 13-27-VII-1902 (ec) [Hdxyyandsm@o1
23-IV-1903 (c). [Have recent records. ]

Amblyscirtes eos (Edwards) 1871. “Comus”. 28-IIJ-1900 (c), 1-IX-1900 (c),
22-11-1901 (s), 11-IV-1901 (ec), 241V-1902 (c), 18-IV-1903 (eB) eee
records, but should be found occasionally. ]

Amblyscirtes celia Skinner 1895. “Celia”. 1-IX-1900 (c), 11-IV-1901 (c).
[Have recent records. |

Euphyes vestris metacomet (Harris) 1862. “Vestris”. 15-IV-1900 (ce); 1-1Xx-
1900 (c). [Have recent records. ]

Atrytone delaware lagus (Edwards) 1881. “H. delaware”. 11-VIII-1901 (c),
21-VIIT-1901 (c-5). [No records since 1901, but should be found rarely. |

Wallengrenia otho egeremet (Seudder) 1863. “Otho”. 1-IX-1900 (c). [Have
recent records. |]

Polites vibex praeceps (Seudder) 1872. “Brettus”. 28-VIII-1907 (c-12),
have not seen it here before. [Have recent records. |
Hesperia viridis (Edwards) 1883. “E. nevada”. 11-II-1901, caught E. nevada
in my hand but let it go as I had nothing to put it in. [Have recent records. |

Hylephila phyleus (Drury) 1773. “Phyleus”. 11-IV-1901 (c). [Have recent

records. |

Copéodes aurantiaca (Hewitson) 1868. “Procris’. 10-IV-1902 (s). [Have

recent records. |

Lerema accius (Smith) 1797. “L. accius”. 3-I-1905 (c), 29-X-1916 (c).

VoLUME 25, NUMBER I 35

Not often seen but has been common at morning glory vines at the house. [Have
recent records. |

Pholisora catullus (Fabricius) 1793. “Catullus”. 6-IV-1900 (s), 18-III-1901 (s),
13-27-VI-1902 (c) [Edwards Co.], 5-XI-1902 (c-2). [Have recent records.]

Celotes nessus (Edwards) 1877. “P. nessus’. 27-III-1900 (c), 30-IV-1901 (c),
18-VI-1902 (c), 13-27-VII-1902 (c) [Edwards Co.]. [Have recent records. ]

Heliopetes domicella (Erichson) 1848. “P. domicella’. 23-VIII-1916 (c).
23-VIII-1916 (c).

[ Stray, no recent records. |

Heliopetes laviana (Hewitson) 1868. “Ericetorum”. 11-VIII-1907 (c). A
day or two ago I got two white hesperids (Ericetorum). [So far as we know Heli-
opetes ericetorum has not been taken in Texas. H. laviana occasionally occurs at
San Antonio, New Braunfels, and Austin. It could very easily range over to Kerr
County in a good year. ]

Pyrgus communis communis (Grote) 1872. “P. tessellata”. 25-II-1900 (c-2),
8-I11-1900 (s), 21-I-1901 (s), 5-XI-1902 (c). [Have recent records. ]

Erynnis brizo (Boisduval & LeConte) 1834. “N. brizo”. 21-II-1900 (c),
8-III-1900 (s), 10-IV-1902 (c). [Burns (1964) located three hybrid examples in
museums from Kerr Co. Texas without dates; these could be them. Burns and the
Kendalls have collected the area without finding E. brizo.]

Erynnis zarucco funeralis (Scudder & Burgess) 1870. “N. funeralis”. 27-IV-
1899 (c-1), 14-III-1900 (c), 13-IV-1900 (s-fresh), 18-V-1901 (c), 5-XI-1902 (c).
[Have recent records. ]

Erynnis martialis (Scudder) 1869. “N. martialis”. 20-III-1899 (c-several),
17-IV-1899 (c), 25-III-1900 (c-1), 13-III-1901 (c-several). [Burns (1964) located
five museum specimens with incomplete dates from Kerr Co. Texas. These dates
should help fill the void. No records since 1901. Careful searching by Burns and
the Kendalls have failed to locate this species in the Kerrville area. ]

Gesta gesta invisus (Butler & Druce) 1872. “Nisoniades llano”. 13-27-VII-1902
(c), fishing trip to Llano River, camped near head of South Llano [River] [Ed-
wards Co.]. 10-31-VIII-1904 (c-1) [Kimble Co.]. [Have recent records. ]

Grais stigmaticus (Mabille) 1883. “Grais stigmaticus”. 20-IV-1903 (c), 30-
VIII-1916 (c), 1-IX-1916 (c-1), 2-IX-1916 (c), 27-IX-1916 (s), 29-X-1916 (c-
several ), 21-IX-1917 (c-5). [Have recent records. ]

Achlyodes thraso tamenund (Edwards) (1807). “Near Grais’. 10-31-VII-1904
(c-l1) [Kimble Co.], 19-VIII-1907 (c-1), 27-IX-1916 (c-1), 29-X-1916 (c1).
[Should be found occasionally. |

Systasea pulverulenta (Felder) 1869. “S. zampa”. 6-IV-1901 (c-1) first I
have ever seen at the ranch. 2-VIII-1901 (s), 8-VIII-1901 (c), 2-VI-1902 (s).
[Have recent records. ]

Cogia hippalus outis (Skinner) 1894. “C. outis’. 23-IV-1899 (c-1), 27-IV-
1899 (c-3), 10-V-1899 (c-2), 7-IV-1900 (s), 8-IV-1900 (c), 15-IV-1900 (c), 6-
IX-1900 (s), 30-III-1901 (s), 4-IV-1902 (s), 18-IV-1902 (s) common now. [Have
recent records. |

Thorybes pylades albosuffusa Freeman 1943. “Pylades”. 17-IV-1899 (c), 1-IV-
1900 (c), 6-IV-1900 (s), 15-IV-1900 (c), 20-III-1901 (c), 7-IV-1902 (c). [Have
recent records. |

Achalarus lyciades (Geyer) 1832. “A. lycidas”. 19-IV-1901 (c), 1-V-1901 (c),
10-V-1901 (c-2), 28-VI-1901 (c). [No records since 1901, probably no longer in the
area. |

Achalarus casica (Herrich-Schaffer) 1869. “Epigena”. 24-VIII-1899 (c), 28-
VIII-1900 (c), 28-IX-1909 (s-2 or 3 lately), 25-IV-1910 (c-2), 26-IV-1910 (c-2).
[No recent records, probably no longer in the area.]

Achalarus toxeus (Plétz) 1882. “A. coyote’. 17-IX-1916 (c). [Should be
found occasionally. ]

36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Autochton cellus (Boisduval & LeConte) 1834. “C. cellus”. 17-IV-1899 (c),
13-IV-1900 (s), 15-IV-1900 (c), 1-IX-1900 (c), 4-IV-1901 (s), 16-IV-1901 (c),
19-IV-1901 (c) commoner than usual, 7-IV-1902 (s). [No recent records, probably
no longer in the area.]

Urbanus proteus (Linnzus) 1758. “E. proteus”. 26-IX-1900 (c), 13-VI-1902
(c), 10-31-VII-1904 (c-1) [Kimble Co.]. [No recent records but should be found
around cultivated beans. |

Urbanus dorantes (Stoll) 1790. “E. rauterbergi”. 23-IX-1900 (c), 11-VIII-
1907 have seen several specimens lately. [No recent records. Should be found
occasionally. ]

Chioides catillus albofasciatus (Hewitson) 1867. “E. albofasciatus”. 24-VIII-
1899 (s), 28-VI-1901 (c), 2-IX-1902 (s), 1-VII-1904 (c-poor specimen), 9-VII-
1904 (s), have seen six specimens since the first. 11-VIII-1907 (c) have taken sev-
eral good specimens lately. [Should be found occasionally. ]

Chioides zilpa (Butler) 1874. “like Albofasciatus”. 1-IX-1916 (c-3), caught
three long-tailed hesperids like albofasciatus, 29-X-1916 (c-1). [Should be found
rarely. |

Epargyreus clarus clarus (Cramer) [1775]. “E. tityrus”. 11-VII-1899 (c),
25-VIII-1899 (c), 11-IV-1901 (s), 9-V-1901 (c), 10-V-1901 (c), 2-VI-1901 (c),
1-VII-104 (c). [Have recent records. ]

Proteides mercurius mercurius (Fabricius) 1787. “Eudamus idas’. 16-VI-
1899 took a large skipper Eudamus idas, have never seen it before. [No records
since 1899, probably a stray. ]

Papilionidae

Battus philenor philenor (Linneus) 1771. “P. philenor’. 30-31-I-1900 (s-
during the month), 3-II-1900 (s), 8-III-1900 (s), 2-III-1901 (c), 5-XI-1902 (s),
12-IX-1916 (saw mantis eating one). [Have recent records.]

Battus polydamas lucayus (Rothschild & Jordan) 1906. “P. polydamas”.
26-IX-1900 (saw one quite close). [Should be found rarely.]

Papilio polyxenes asterius Stoll 1782. “P. asterius’. 19-III-1899 (s), 8-III-
1900 (s), 8-IV-1900 (c), 2-III-1901 (c). [Have recent records. |

Papilio cresphontes cresphontes Cramer [1777]. “P. cresphontes’. 21-III-1899
(s), 24-IIT-1900 (s), 1-IV-1901 (s). [Have recent records. ]

Papilio glaucus glaucus Linnzeus 1758. “P. turnus’. 19-III-1899 (s), 9-III-
1900 (s), 17-III-1901 (s), 13-27-VII-1902 (c) [Edwards Co.], 29-II-1908 (s).
[Probably western distribution limit. Should be found occasionally. ]

Papilio multicaudata Kirby 1884. “P. daunus”. 29-VIII-1899 (c) Kerrville,
6-IX-1900 (s), 17-VIII-1902 (c), 18-VIII-1902 (c), 5-IX-1908 (saw several times
lately), 15-IV-1910 (c-2) it is not as a rule common but it is by no means rare
some years, 18-IX-1912 (c). [Have recent records. |

Papilio troilus Linnzeus 1758. “P. troilus’. 11-III-1900 (s), 14-III-1901 (c-
fresh), 17-IX-1903 (found one caught by a green-grey spider). [Western distribu-
tion limit. Have recent records.]

Pieridae

Pieris protodice protodice Boisduval & LeConte 1829. “P. protodice”. 12-II-
1900 (s), 8-III-1900 (s), 4-III-1901 (s), 5-XI-1902 (c). [Have recent records.]

Ascia monuste phileta (Fabricius) 1775. “P. monuste”. 11-VIII-1901 (c),
L1-VIiI-1907 (c) commoner than usual, 14-VIII-1914 (c) fairly common for the
last week or two. [Should be found occasionally. ]

Colias eurytheme eurytheme Boisduval 1852. “C. eurytheme” also “Ariadne”.
21-IJ-1899 (c-several), 22-II-1899 (s), 3-II-1900 (c), 21-II-1900 (c-5), 8-III-1900

VOLUME 25, NUMBER 1 37

(s), 8-IV-100 (have seen several winter forms lately), 18-I-1901 (c-1) white form,
31-I-1907 (s), 24-I-1909 (s). [Have recent records. ]

Colias cesonia (Stoll) 1790. “C. cesonia’. 22-II-1899 (s), 8-III-1900 (s),
18-II-1901 (saw several), 7-III-1909 (c). [Have recent records. ]

Anteos clorinde nivifera Fruhstorfer 1907. “Gonepteryx clorinde”. 25-IX-1900
(c), 26-IX-1900 (c), 27-IX-1900 (s), 10-VI-1901 (s), 2-IX-1916 (c-1), (s-2), 8-
IX-1961 (c), 29-X-1916 (c-2). [Should be found occasionally. ]

Phoebis senne eubule (Linnzus) 1767. “C. eubule”’. 8-!II
1902 (c) quite common this month, 5-XI-1902 (s), 28-I-1903 (
records. |

Phoebis agarithe maxima (Neumoegen) 1891. “C. agarithe”. 3-VI-1899 (c-2),
15-VI-1899 (c), 22-VHII-1901 (s), 3IV-1902 (s), 7-IV-1902 (s), 11-VIII-1907
(fairly common, hard to get perfect), 23-VIII-i916 (c) very common, 29-X-1916
(has been very common). [Have recent records. ]

Kricogonia lyside (Godart) 1819. “K. lyside”. 10-V-1899 (c-5), 12-V-1899
(common), 12-IV-1900 (s), 7-XI-1900 (scarce), 9-IV-1902 (s) quite a number,
all flying in southwesterly direction singly; wind south, 10-IV-1902 (c), 14-IV-1902
(of the hundreds that crossed the road as I was riding to town Saturday, everyone
of them was going in a southerly direction), 30-IV-1902 (out fresh), 18-VI-1902
(c), 27-V-1903 (c-fresh), 29-VIII-1907 (very common), 1-VI-1918 (swarms, 3 or
4 forms, were passing the ranch today from soon after noon till about 4 p.m., all
going East. Lena Denton phoned from the Masterson ranch to say the same thing
was happening there. There was a constant stream of them crossing the trail. 22-
VI-1918 (still numerous), 11-IV-1919 (s) few, 12-IV-1919 (going south), 13-IV-
1919 (still going south). [Have recent records. ]

Eurema mexicana (Boisduval) 1836. “E. mexicana’. 30-IV-1900 (c) numerous.
[ Have recent records. |

Eurema lisa Boisduval & LeConte 1829. “E. lisa’. 21-I-1901 (s), 5-XI-1902
(s), 26-I-1909 (c). [Have recent records. ]

Eurema nicippe (Cramer) [1780]. “Nicippe”. 21-I-1901 (s), 5-XI-1902 (s).
[Have recent records. |

Nathalis iole Boisduval 1836. “N. iole’. 22-II-1899 (s), 30-31-I-1900 (s)
this month, 8-III-1900 (s), 18-I-1901 (c), 21-I-1901 (c), 5-XI-1902 (c), 24-I-
1909 (s). [Have recent records. ]

Anthocharis midea midea (Hiibner) [1809]. “Genutia”. 15-IV-1900 (c), 11-
1V-1901 (ec). 7-IV-1902 (c), 16-IV-1902 (c), 1-III-1909 (s), 7-III-1909 (c).
[Have recent records. |

1900) (s)e axe
s). [Have recent

Riodinidae

Calephelis australis Edwards 1877. “C. australis”. 1-IX-1900 (c), 18-IV-1901
(c), 18-27-VII-1902 (c) [Edwards Co.]. [Have recent records.]

Calephelis rawsoni McAlpine 1939. Not mentioned by Lacey. Holotype male
and one paratype male collected by Lacey, July 1908 Kerrville, Texas and just Kerr-
ville, Texas respectively. It is unlikely that Lacey actually collected these speci-
mens at Kerrville. The authors have reared this species from vicinity of Lacey’s old
ranch but have been unable to locate suitable habitats in or very near Kerrville.

Lycaenidae

Harkenclenus titus watsoni (Barnes & Benjamin) 1926. “T. titus’. 11-VI-
1900 (c), 21-VI-1900 (c) on button willows [Cephalanthus occidentalis] which are
just out, 17-VI-1901 (c-2), 25-V-1902 (c-1?). [No recent records but should be
found occasionally. |

Satyrium calanus falacer (Godart) [1824]. “T. calanus”. 12-V-1899 (c), 17-V-

38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

1899 (c-20) all fresh, 15-V-1900 (c), 14-V-1901 (c), 18-V-1901 (c) about 20, 14-
V-1902 (c) first few I get are always darker than the later ones, 21-V-1902 (com-
mon), 24-V-1903 (c). [Have recent records. |

Calycopis beon (Stoll) 1782. “T. cecrops’. 27-III-1900 (s), 2-III-1901 (c),
13-VI-1902 (s), 18-VI-1902 (c). [Have recent records.]

Tmolus azia (Hewitson) 1873. Not listed in Lacey’s field notes. Recorded by
Barnes & McDunnough (1913). Collected by Lacey (19), Paint Creek, Edwards
Co. Texas. No date given. According to Lacey’s field notes, he visited Paint Creek
15 July 1898, 23 November to 6 December 1916, and 19-26 July 1919; the last two
dates are ruled out for obvious reasons, leaving 15 July 1898 most likely date of
capture. [A stray. ]

Callophrys henrici solatus Cook & Watson 1909. “I. irus”. 20-III-1899 (c-6),
21-III-1899 (c-4), 8-III-1900 (c), 13-III-1901 (c-3), 15-III-1901 (c) several, 16-
III-1903 (c). [Single brooded. Have recent records. ]

Callophrys gryneus castalis (Edwards) 1871. “T. damon”. 13-III-1899 (c),
21-III-1899 (c-2) [Kimble Co.], 26-VII—5-VIII-1899 (c) several very dark, 9-III-
1900 (s), 2-III-1901 (c), 30-IV-1902 (c). [Have recent records. ]

Atlides halesus corcorani Clench 1942. “A. halesus”. 15-VI-1899 (c-2), 25-
VIII-1899 (c), 27-III-1900 (s), 1-IX-1900 (c), 21-VIII-1901 (s), 13-VI-1902 (s),
18-VI-1902 (c), 7-III-1909 (c). [Have recent records. ]

Euristrymon ontario autolycus (Edwards) 1871. “Autolycus”. 15-V-1900 (c),
17-V-1901 (c) 40 good specimens, 21-V-1902 (common), 24-V-1903 (c). [Have
recent records. |

Panthiades m-album (Boisduval & LeConte) [1833]. “T. m-album”. 29-VIII-
1907 (c) first I have ever seen. [No records since 1907. Probably western distribu-
tion limit. ]

Strymon melinus franki Field 1938. “T. melinus”. 14-III-1900 (c), 24-V-
1903 (c) on milkweed blossoms. [Have recent records. |

Strymon columella istapa (Reakirt) [1867]. “T. columella”. 25-VI-1902 (c)
first I have seen here. [Should be found occasionally. Have one recent record.]

Strymon alea (Godman & Salvin) [1887]. Callicista laceyi Barnes & McDun-
nough 1910. Not mentioned in Lacey’s field notes. C. laceyi was described from a
female collected by Lacey in July 1909 at Del Rio, Texas. Lacey left Kerrville 15
July 1909 for the Devils River on a fishing trip. He stayed until 31 July. It is un-
likely that he passed through Del Rio which would have been considerably out of
his way. The most likely place of capture was on the Devils River north of Del
Rio in Val Verde Co.

Leptotes marina (Reakirt) 1868. “L. marina’. 10-V-1899 (c-1). [Should be
found there occasionally. |

Hemiargus ceraunus zach@ina (Butler & Druce) 1872. “L. gyas”. 5-XI-1902
(c-1), like isola without black spots beneath upper wing, 29-X-1916 (fairly com-
mon). [Have recent records. ]

Hemiargus isola alee (Edwards) 1871. “Isola”. 8-IV-1900 (s), 5-XI-1902 (c),
10-II-1903 (c). [Have recent records. ]

Everes comyntas comyntas (Godart) [1824]. “E. comyntas’. 27-IV-1899
(c-19), 9-VIII-1899 (c), 24-III-1900 (s), 21-VI-1900- (c), 30-III-1901 (c) have
been out a few days. [No recent records, but should be found occasionally. ]

Libytheidae

Libytheana bachmanii larvata (Strecker) [1878]. “L. bachmanii’. 7-XI-1900
(scarce), 24-IV-1901 (c), 5-XI-1902 (c), 28-I-1903 (s), 10-II-1903 (s) out every
rm day throughout the winter, 13-VIII-1916 (swarming), 20-VIII-1916 migrat-

% in countless thousands all going southeast, 21-VIII-1916 still going east in less
12-IX-1916 hundreds of thousands: on the verbesina weeds [Verbesina

VOLUME 25, NuMBER l 39

virginica L.] last few days, 25-IX-1916 still going southeast, 5-X-1916 large flight
went by the ranch, 8-X-1916 still migrating, 16-X-1916 few disreputable specimens
going east, 29-X-1916 swarming and migrating. [Have recent records. ]

Libytheana carinenta mexicana Michener 1943. “Carinenta’. 5-XI-1902 (c).
[Probably a stray. ]

Nymphalidae

Anaea andria andria Seudder 1875. “A. andria’. 18-III-1900 (c), 13-III-1901
(s), 7-III-1909 (c), 4-I-1917 (c). [Have recent records. ]

Anaea aidea morrisonii (Edwards) 1883. “A. morrisonii”. 7-XI-1900 (c).
[Should be found rarely. ]

Mestra amymone (Ménétriés) 1857. “C. amymone’. 8-IX-1898 (c-1), 27-IX-
1898 (c-1), 18-VI-1900 (s), 22-VI-1900 (s), 7-VII-1900 (c) [Kimble Co.], 25-VI-
1902 (c). [Have recent records. |

Marpesia chiron (Fabricius) 1775. “Timetes chiron”. 28-VIII-1899 (c-1) have
been looking for him every day since I saw him on the 24th, 3-VIII-1902 (s), 14-
VII-1905 (s-worn), 11-VIII-1907 (c-1) saw another, 25-VIII-1907 (s) third seen
this year, 1-IX-1916 (s-1). [Should be found rarely. ]

Marpesia petreus (Cramer) [1776]. “T. peleus’. 14-VIII-1907 (c-2), 29-VIII-
1907 (c-1) a good specimen, third taken and fourth seen this year. [Should be
found rarely. |

Limenitis astyanax astyanax (Fabricius) 1775. “Ursula”. 30-IV-1902 (c) they
have been out for about a week. [Have recent records. ]

Limenitis archippus watsoni (dos Passos) 1938. “L. disippus” [Possibly hy-
brid]. 25-VI-1899 (c) at San Antonio [Bexar Co. Texas], 14-VII-1905 (c-1) fresh,
it is not at all common here. [Have recent records. |

Limenitis bredowii eulalia (Doubleday) [1848]. “H. californica’. 22-VIII-
1900 (s-1) saw another about two years ago, 29-IX-1900 (s-2), 3-XI-1900 (s-1),
4-XT-1900 (s), 13-27-VII-1902 (c) [Edwards Co.], 8-VIII-1903 (s), 17-IX-1903 (c),
18-IX-1903 (s), 29-IX-1903 (s-2) fresh, 30-IX-1903 (s), 15-IV-1904 (s-3), 18-
VI-1904 (c), 9-VII-1904 (s), 29-VIII-1907 (s) fresh, 12-IV-1910 (s-several) fresh,
3-IV-1916 (s) fresh, 29-X-1916 (fairly common). [Occasionally becomes temporary
resident. Have recent records. |

Metamorpha steneles biplagiata (Fruhstorfer) 1907. “V. steneles”. 26-27-VI-
1904, saw what I suppose was V. steneles but the ground color was yellowish in-
stead of white; tried hard to get him but failed. [Should stray into area rarely.]

Vanessa atalanta (Linneus) 1758. “Atalanta”. 30-I-1900 saw this month, 4-II-
1900 (s), 8-III-1900 (s), 21-I-1901 (s), 5-IX-1902 (s), 24-I-1909 (s), 4-I-1917
(s). [Have recent records. |

Vanessa virginiensis (Drury) [1773]. “Huntera”’. 30-I-1900 saw this month,
8-III-1900 (s), 21-I-1901 (s), 17-II-1901 (s) often very small in winter and early
spring [attributed to scarcity of larval food plant], 5-XI-1902 (c). [Have recent rec-
ords. |

Vanessa cardui (Linneus) 1758. “P. cardui’. 8-III-1900 (s), 21-VIII-1901
(c) common, 16-XII-1902 (s). [Have recent records. ]

Junonia coenia coenia (Hubner) [1822]. “J. coenia’. 20-III-1900 (c), 13-IV-
1900 (s), 5-XI-1902 (s), 26-I-1909 (c). [Have recent records. ]

Nymphalis antiopa lintnerii (Fitch) 1856. “V. antiopa’. 25-II-1900 (s)
worn, 17-V-1900 (s), 31-V-1900 (c), 18-II-1901 (s), 2-III-1901 (s), 8-V-1902 (c),
97-V-1903 (s-2), 28-V-1903 (c-1), 31-V-1903 (c-1), 22-II-1907 (s) more than
ever before, 7-V-1911 (c-several); not common at the ranch. Saw none in 1899
and only two in 1898. [Should be found occasionally. ]

Polygonia interrogationis (Fabricius) 1798. “C. interrogationis’. 22-II-1899

>

AQ JouRNAL OF THE LEPIDOPTERISTS SOCIETY

(s), 4-II-1900 (s), 30-IV-1900 (c) numerous, fresh, 30-IV-1902 (c). [Have recent
records. |

Polygonia comma (Harris) 1842. “P. comma”. 15-IV-1900 (s), 11-IV-1901
(c). [No records since 1901. Probably no longer present. |

Chlosyne janais (Drury) 1782. “Synchloe janais”. 27-VI-1899 (c) [Bandera
Co.], 22-VII-1908 (s) in garden. [Should stray into the area occasionally. ]

Chlosyne lacinia adjutrix Scudder 1875. “S. lacinia”. 27-VI-1899 (c) [Bandera
Co.], 13-IV-1900 (s), 5-IV-1901 (c), 15-IV-1902 (s), 16-IV-1902 (c), 5-XI-1902
(c), 5-XI-1902 (s), 11-VIII-1903 (1-ex larva; larva found on white ragweed
[Parthenium hysterophorus L.], 31-VIII-1903 (1-ex larva). [Have recent records.]

Chlosyne endeis (Godman & Salvin) 1894. Not listed in Lacey’s field notes.
Recorded by Barnes & McDunnough (1913). Collected by Lacey (1¢, 12); 6
marked “Texas,” the @ May 1902, Edwards Co., much worn. [Strays].

Phyciodes texana texana (Edwards) 1863. “E. texana’. 21-I-1901 (s), 2-III-
1901 (s), 30-IV-1902 (s) fresh, 25-VI-1902 (c), 20-II-1909 (c). [Have recent rec-
ords. |

Chlosyne nycteis nycteis (Doubleday) [1847]. “P. nycteis”. 12-IV-1900 (s),
30-III-1901 (s), 10-IV-1902 (s), 16-IV-1902 (c). [Have recent records. ]

Chlosyne gorgone carlota (Reakirt) 1866. “Ph. ismeria’. 19-IV-1899 (c).
[No recent records, but should be present. ]

Phyciodes tharos tharos (Drury) [1773]. “Ph. tharos”. 13-III-1900 (s), 18-
III-1900 (s), 7-III-1909 (c). [Have recent records. |

Phyciodes phaon (Edwards) 1864. “Phaon”. 18-III-1900 (c), 30-III-1901 (c),
5-XI-1902 (s), 7-IJI-1909 (c). [Have recent records. |

Phyciodes vesta (Edwards) 1869. “Ph. vesta”. 2I1-III-1899 (s), 18-III-1900
(c), 18-III-1901 (s), 25-IV-1901 (c), 7-IV-1902 (c), 10-IV-1902 (c), 5-XI-1902
(c), 24-I-1909 (s). [Have recent records. ]

Thessalia theona bollii (Edwards) [1878]. “M. bollii”. 25-V-1900 (c) first I
have ever seen, 28-V-1906 (c-1), 21-VI-1900 (c-3), 6-IX-1900 (s), 19-V-1908 first
I have seen here for years, 5-IX-1908 (c) fairly common on the hill, 7-IX-1908 (c)
more on the hill. [Have recent records. |

Dymasia dymas (Edwards) 1864. “M. dymas”. 25-26-VI-1899 (c) San Antonio
[Bexar Co.]. [Have recent records, but urbanization is rapidly destroying habitats. ]

Texola elada ulrica (Edwards) 1877. “M. elada’. 15-26-VII-1898 took good
series [Edwards Co.], 25-30-VII-1901 (c) [Kimble Co.], 8-VIII-1901 (c), 30-IV-
1902 (c). [Have recent records. |

Poladryas minuta minuta (Edwards) 1861. “M. arachne’. 15-VII-1898 (c-2)
[Edwards Co.], 27-IX-1898 (c-18) near [ranch] gate, 21-VI-1900 (c), 14-IV-1901
(s), 20-IV-1901 (c-45), 21-VIII-1901 (s), 22-IV-1902 (s) several, 23-IV-1902 (c),
30-IV-1902 (common), 9-VI-1902 (s) many on Kerrville road, 13-VI-1902 (fairly
common), 21-VIII-1904 fresh brood out, 7-IX-1908 (c-2), 26-I-1909 (s) not seen
generally until April, 15-VIII-1909 (c) several, 12-IV-1910 (s), 14-II-1911 (s) along
Turtle Creek road. [No records since 1911. Habitats probably destroyed by over
grazing of sheep and goats.]

Euptoieta claudia (Cramer) [1776]. “E. claudia”. 31-I-1900 (s) this month,
4-II-1900 (s), 21-II-1900 (c), 7-XI-1900 (c), 17-11-1901 (Ss); 52 xqeisoz eee
| Have recent records. |

Euptoieta hegesia hoffmanni Comstock 1944. “E. hegesia”. 15-VIII-1909 saw

fine fresh E. hegesia but failed to get it. [Should stray into the area rarely. |]
Heliconius charitonius vazqueze Comstock & Brown 1950. “Heliconius
charitonius.” 26-VII-1898, one of the Miller boys caught it by hand in Kerrville and
brought it to Palmer’s place. I have not seen it here before. 19-VII-1908, saw one
in the garden but couldn’t get it. [Migrant. Should be found there occasionally. |

Dryas julia moderata (Stichel) 1907. “Coleenis julia”. 7-X-1900, Jim Taylor

VoLUME 25, NuMBER 1 Al

took one last month and brought it to me to identify. 6-X-1906 (s-2), 11-VIII-1907
(c-1) at the ranch; better than I got in Mexico. 14-VIII-1907 (c-1) saw three
others. [Migrant. Should be found occasionally. ]

Agraulis vanillae incarnata (Riley) 1926. “A. vanille”. 24-IV-1900 (s)
fresh, 2-II-1901 one settled three times on the stem of my pipe as I was having a
smoke near the little dam, 5-XI-1902 (s) fresh. [Have recent records. ]

Danaidae

Danaus plexippus plexippus (Linnzus) 1758. “D. archippus”. 28-X-1882
crowds going south, 23-X-1892 were passing over all day, 25-III-1900 common,
weary looking, 17-V-1900 (s) fresh, 13-III-1901 (s) worn, 30-IV-1902 (c) fresh,
5-XI-1902 (s) fresh, 27-V-1903 (s-1) fresh, 19-X-1904 in swarms, 29-II-1908 (s)
fresh, 8-X-1916 gathering about the willows and pecans, 10-X-1916 going south in
hundreds, 31-III-1917 going north, not in the big bunches that go south in the fall,
5-X-1918 for the last week they have been swarming in the pecan.

Danaus gilippus strigosus (Bates) 1864. “Strigosus”. 21-I-1901 (s), 13-III-
1901 (s), 5-XI-1902 (s), 24-I-1909 (s). [Have recent records. ]

Satyridae

Euptychia hermes sosybius (Fabricius) 1793. “N. sosybius”. 11-VI-1900 (c),
1-VII-1904 (s). [Should be found occasionally. ]

Euptychia cymela cymela (Cramer) [1777]. “Eurytus’. 15-V-1900 (c). [Possi-
bly western distributional limit. Have recent records. ]

Euptychia rubricata Edwards [1871]. “N. rubricata”. 5-V-1900 (s), 28-VIII-
1900 (c), 6-IX-1900 (c), 4-IIIJ-1901 (s-1), 6-IV-1901 (c), 16-IV-1901 (c), 8-V-
1902 (c). [Have recent records. ]

Cercyonis pegala texana (Edwards) 1880. “S. texana”’. 21-VI-1900 (c),
7-19-VII-1900 (c) [Kimble Co.], 16-VI-1901 (s) several, 9-VI-1902 (s). [Have
recent records. |

Sphingidae

Manduca sexta (Johanssen) 1763. “Carolina”. 12-IX-1916 yesterday I took
a large Carolina sphinx from a mantis. [Have recent records. ]

Sphinx eremitoides Strecker 1874. “H. eremitoides’. 12-IX-1916 a mantis was
eating an H. eremitoides, 13-IX-1918 yesterday I noticed 3 or 4 H. eremitoides rest-
ing on posts while fixing fence. This morning I went around with a poison jar and
gathered in about 20. Their color matched the cedar posts exactly. 14-IX-1918
(c-6), 22-IV-1919 many on fence posts, 23-IV-1919 more, quite a bunch of them.
[Have recent records, but never found common. ]

Hemaris thysbe (Fabricius) 1775. “Thysbe”’. 10-III-1900 (c), 25-III-1900
watched female laying its eggs on flower buds of black haw [Crataegus tracyi
Ashe]. Last year I found the caterpillars of this moth feeding on honeysuckle
[Lonicera albiflora T. & G.] 13-III-1901 (c-1). [Should be found occasionally.]

Hemaris diffinis (Boisduval) 1836. “H. axillaris”. 10-III-1900 (c). [Should
be found occasionally. ]

Amphion nessus (Cramer) 1777. “Amphion nessus” 28-IIJ-1900 (c). [Should
be found occasionally. ]

Xylophanes tersa (Linnezus) 1771. 7-XI-1896 on Nueces [River] [Uvalde Co.]
picked up a sphinx larva with false eyes on it like the elephant hawk at home
| Deilephila elpenor L.| [Have recent records. |

Hyles lineata (Fabricius) 1775. “C. lineata’. 10-III-1900 (c-2), 19-V-1908
screech owl feeding her young Sphinx moth (lineata). [Have recent records. ]

49, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Saturniidae

Hyalophora cecropia (Linnzus) 1758. “Cecropia’. 6-IV-1900 (1) ex pupa,
another 3 or 4 days ago. 3-IV-1901 (1) ex pupa, 12-V-1901 (1) ex pupa, 24-V-
1903 (1) ex pupa. [Should be found occasionally. ]

Actias luna (Linnzus) 1758. “Luna”. 6-III-1908 yesterday I saw a beautiful
fresh luna moth. [Have recent records. ]

Antheraea polyphemus (Cramer) 1775. “T. polyphemus”. 11-VI-1902 on the
first of the month Rawson got me a female T. polyphemus; she laid a lot of eggs
which I brought home and the larvae were hatched this morning. 6-III-1918 (c).
[Have recent records. ]

Hemileuca maia (Drury) 1773. “Hemileuca maia”’. 4-XI-1900 (c-1) first this
year, 11-XI-1902 (1) ex larva, 18-XI-1902 a female hatched from one of my pupae.
I took it outside with a bit of mosquito curtain over the box. It was immediately
surrounded by males. All the males came up wind to the box and when they over
ran the scent, circled back to it like a pack of hounds. 31-X-1903 (s), 23-X-1904
(c), 11-XI-1912 (s) first this season. [Have recent records.]

Hemileuca grotei Grote & Robinson 1868. “H. grotei’. 5-XI-1902 (c), 7-XI-
1902 (192) ex larva, 9-X-1904 (19) ex larva. [Have recent records. ]

Eacles imperialis (Drury) 1770. “Eacles imperialis”. 26-VIII-1901 (c). [No
recent records. Probably western distribution limit. ]

Amatidae

Horama texana Grote 1867. “Horama texana’”. 1-IX-1916 (c) first I have
seen here, 2-IX-1916 (c). [No recent records but should be found occasionally.]

Arctiidae

Apantesis arge (Drury) 1770. “Arge”. 7-XI-1900 (c), 26-II-1901 watched
arge deposit eggs on a dead grass stem. [Have recent records.]

Hyphantria cunea (Drury) 1770. “H. cunea”. 16-X-1900 caterpillars have
stripped most of the pecans and walnuts of their leaves; ranchers are busy attending
to the caterpillars. 6-X-1909 worse than ever, the caterpillars get into everything
and in some places are all over the ground. 10-X-1916 webworms (H. cunea) very
numerous this year. [Have recent records. ]

Noctuidae

Pseudaletia unipuncta (Haworth) 1809. “L. unipuncta”. 9-X-1904 very
common a week or two ago, 26-XI-1904 numerous caterpillars this year. [Have re-
cent records. |

Lithophane laceyi (Barnes & McDunnough) 1910. Not mentioned by Lacey
in his field notes. Described from specimens collected by Lacey. Should be there.

Spodoptera frugiperda (Smith) 1797. “L. frugiperda”. 9-X-1904 very common
a week or two ago, 26-XI-1904 same as above. [Have no recent records. |

Catocala consors (Smith) 1797. “Consors”. 25-V-1902 (c) out now. [Have
recent records. |

Sepa junctura Walker 1857. “Junctura”. 25-V-1902 (c). [Have recent
records.

Catocala frederici Grote 1872. “Catocala frederici”. 31-V-1903 (c) at night.

This is the fourth of this rare species that I have taken. [Should be found rarely. |
eatin ultronia (Hiibner) 1823. “Ultronia”. 25-V-1902 (c). [Should be
taken rarely, |

Catocala similis Edwards 1864. “Similis”. 95-V-1902 (c). [Should be found

alu |
iy

VoLUME 25, NuMBER 1 43

Catocala micronympha Guenée 1852. “Micronympha”. 25-V-1902 (c). [Have
recent records. |

Catocala amica (Hubner) 1818. “Amica”. 25-V-1902 (c). [Have recent
records. ]

Alabama argillacea (Hiibner) 1823. “A. argillacea”. 9-X-1904 swarms: every-
where, 26-XI-1904 numerous. [No recent records but should be found regularly. ]

Notodontidae

Datana ministra (Drury) 1773. “Caterpillars”. 27-VI-1899 the pecan trees
on the Medina [River] [Bandera Co.] are completely stripped of their leaves by
caterpillars, something like those of our buff-tip moth [Phalera bucephala L.] [Have
recent records. |

Lasiocampidae

Malacosoma disstria Hiibner 1820. “M. disstria”. 23-IV-1900 last Friday and
Saturday the train was delayed for half an hour in the neighborhood of Boerne,
Texas [Kendall Co.] by the quantity of caterpillars on the rails that prevented the
wheels from getting a grip on them. These caterpillars are completely stripping the
oak trees. [This species was very abundant during 1959-1964 throughout central
Texas. It has since been under control, apparently through natural biological means.]

Megalopygidae

Lagoa laceyi Barnes & McDunnough 1910. Not mentioned by Lacey. De-
scribed from specimens collected by Lacey. No recent records, but it should be found
there rarely.

Acknowledgment

This article is dedicated to the memory of our very dear friend and
fellow Lepidopterist, Mrs. Ellen Schulz Quillin who died suddenly at her
home in San Antonio, Texas 6 May 1970. Mrs. Quillin was founder of the
Witte Memorial Museum and Director for thirty-seven years, and Direc-
tor Emeritus until her death. She was a noted botanical author and lec-
turer. Without her foresight in acquiring the hand scribed field notes
of Howard George Lacey, these data could not now be made available
to the Lepidopterists’ community.

Literature Cited

BARNES, WILLIAM, AND J. McDuNNoucH. 1910. A new Thecla from Texas. Can.
Ent. 42: 365-366.

1913. Species of Lepidoptera new to our fauna, with synonymical notes.
Can. Ent. 45: 182-185.

Burns, Jonn M. 1964. Evolution in skipper butterflies of the genus Erynnis.
Univ. Calif. Pub. in Ent. 37: 44-64.

Cuiencu, H. K. 1966. The synonymy and systematic position of some Texas
Lycaenidae. J. Lepid. Soc. 20: 65-66.

pos Passos, Cyrm F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid.
Soc. Mem. 1.

44 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

1969. A revised synonymic list of the Nearctic Melitaeinae with taxonomic

notes (Nymphalidae). J. Lepid. Soc. 23: 115-125.

1970. A revised synonymic catalog with taxonomic notes on some Nearctic
Lycaenidae. J. Lepid. Soc. 24: 26—38.

McAtping, W. S., 1939. A new Metal Mark (Calephelis) from Texas. Bull. Brook-
lyn Ent. Soc. 34: 75-80.

McDunnovucu, J. 1938. Checklist of the Lepidoptera of Canada and the United
States of America, Part 1, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1 (1).

1939. Checklist of the Lepidoptera of Canada and the United States of

America, Part 2, Microlepidoptera. Mem. So. Calif. Acad. Sci. 2 (1).

A REVISION OF SPEYERIA NOKOMIS (NYMPHALIDAE)?*

C.LirFoRD D. FERRIS
University of Wyoming, Laramie, Wyoming

MikE FISHER
1200 Summit Road, Ponderosa Hills, Parker, Colorado

It is proposed that Speyeria nokomis nitocris (Edwards) be elevated
from the synonymy as placed by dos Passos (1964), to subspecific rank
for the reasons which are delineated in the paragraphs below.

In an earlier work (dos Passos and Grey 1947), Speyeria nokomis
nitocris was treated as a valid subspecific taxon, but later (1964), dos
Passos treated this taxon as synonymous with S. nokomis nokomis (Ed-
wards). For the past several years, we have made a study of this insect
in Colorado (M. F.) and in Arizona and New Mexico (C. D. F.). The
Colorado insect, S. nokomis nokomis is quite distinct in habitat and fascia
from the Arizona-New Mexico insect, S. nokomis nitocris.

Speyeria nokomis nokomis (Edwards )

Original Description: “Descriptions of certain species of diurnal Lepidoptera found
within the limits of the United States and British America.” No. 3. Proc. Acad.
Nat. Sci. Philadelphia 14: 221 (1863). Figured by Edwards, B. N. A. pl.
Argynnis 4 (1868).

Type Locality: This was fixed by dos Passos and Grey (1947) as Mount Sneffels,
Ouray Co., Colorado. This location has been questioned by Brown (1957b, p. 335)
and does not appear consistent with presently known habitats for n. nokomis. The
neotype male (AMNH) bears the label “Oslar Sneffels Mts Ouray Co Col Aug
9000 Ft.,” “A. nokomis.” and “Ex Coll. Wm. C. Wood Acc. 36915.”

Discussion: In the plate which accompanies this paper, three sub-
species of Speyeria nokomis are figured. Dorsally the males are similar

1 Published with the a

Journal Paper No. 434. pproval of the Director, Wyoming Agricultural Experiment Station, as

Votume 25, Numser | 45

but distinct differences are evident on the ventral surfaces. In S. n. no-
komis, the discal area of the hind wing is a pale cimnamon brown. The
silver spots are large, brilliant, and moderately edged with black. The
species as a whole is strongly dimorphic (excepting n. wenona) and the
females dorsally tend almost to black basally. In n. nokomis, dorsally the
females show a blue-green overscaling in the dark areas with a light buff
submarginal band. On the primary, the band shows a yellow-orange
flush; on the secondary the flush is bluish. Ventrally, the females are
similar to the males as far as the primaries are concerned, except that the
colors are more intense and the ground color is a darker red. On the
secondary, the submarginal band is a light buff and the ground color
of the discal area is buff overscaled with a greenish-black dusting. Fe-
males from the Paradox Valley colony in Colorado tend to be more blue
dorsally than specimens from other localities.

This insect is currently known from several locations in Colorado, from
Uintah Co. in eastern Utah, and in a blend with n. apacheana from more
westerly areas of Utah. The Colorado locations are Unaweep Canyon,
Mesa Co., 6000’ and Paradox Valley, Montrose Co., 5200’. The Colorado
habitats are nearly neutral seep areas where there is a constant flow of
water and are surrounded by willow thickets. The violets which serve as
the larval foodplant grow in the understory of the seeps and are found
only sparsely in the thickets. The surrounding region, except in the vi-
cinity of other seeps and washes is relatively arid. The sites of both colo-
nies are unusual with respect to their geology. The usual rock formation
in this portion of Colorado is Mancos shale. The Unaweep Canyon seep
emerges along a spring line in granite and flows over what appears to be
stream alluvium. The canyon is unusual in that it is mainly composed of
granite. The Paradox Valley colony is in a blowsand area. The Uintah
Co., Utah area is a rather unrewarding looking meadow, but with the
necessary prerequisites of willow, violets, and a constant flow of water.
The presence of willow is probably only incidental to the presence of
nokomis. Both willows and violets are frequently found in the same loca-
tion in arid regions.

There is one additional recent record of S. nokomis from Colorado.
This is a worn female taken by Scott Ellis at Rogers Mesa Delta Co., 5850/
26 August 1958. The source of this specimen has not yet been determined.
although it perhaps came from an undiscovered Colorado colony.

Dos Passos and Grey (1947) fixed the type locality for S. nokomis no-
komis as Mount Sneffels, Ouray Co., Colorado. We feel that we must
take exception to this fixation for several reasons. Collecting im recent

46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

years in this area (Brown, in litt.) has failed to turn up this butterfly.
The ecology of the fixed type locality is not consistent with currently
known collection areas. Furthermore the neotype does not fit the original
description (Brown, 1965). There is also a disparity in dates. Dos Passos
(1964) indicates an 1862 (“1861-3”) date for the Edwards publication
of the description of n. nokomis. If this is the correct date, it would not
seem possible for the specimens to have come from Ouray Co., as the
first whites to enter this area came in 1871. Oslar is known to have been
quite inaccurate in his data. As Brown has mentioned (1965), the Ma-
comb Expedition entered the area west of Paradox Valley in 1859 (Ma-
comb and Newberry 1876). It is possible that the material ascribed to
Edwards 1862 was actually collected on this expedition. Brown placed
the probable type locality for S. nokomis nokomis (Edwards) (and not
for the neotype S. nokomis nokomis dos Passos and Grey) as Ojo Verde,
San Juan Co., Utah. The location Ojo Verde is mentioned in the Ma-
comb/Newberry report. The topographical coordinates are given in the
report as lat. 38° 14’ 50” and long. 109° 26’ 40’. Present United States
Coast and Geodetic Survey maps do not show Ojo Verde, but the co-
ordinates would indicate it to be in the vicinity of Hatch Wash. In Au-
gust, 1969, the senior author attempted to locate this site. He was un-
successful as that portion of Utah is now posted as an impact area for
test missiles launched from Green River, Utah. The warning notices and
travel directions were sufficiently ambiguous that he did not wish to risk
entry into the area.

The comments in the Macomb/Newberry report would appear to in-
dicate a possible habitat for n. nokomis. Macomb and his associates
camped at Ojo Verde in mid-August and would thus have been there at
the correct time of the year for the insect’s flight. Of the area, the fol-
lowing comments are made: “The Ojo Verde is a copious spring in a
canon cut out of the red sandstone, ten miles west of La Teneja. The
surrounding country is very sterile, sparsely set with sage bushes and
small cedars, but about the spring the bottom of the cafion is covered
with the greenest and most luxuriant grass.”

It would appear then that the type locality for S. nokomis nokomis
(Edwards) is most probably somewhere in eastern Utah, and not in
Ouray Co., Colorado, although L. P. Grey now favors the Rio Hondo
area in Lincoln Co., New Mexico.

There also exists a record of a single male and two females of S. n.
nokomis taken September 15, 1877 by Lt. C. A. H. McCauley at the cross-

of the Lower Rio Florida, La Plata, Co., Colorado 6500-7000’ some

VOLUME 25, NuMBER 1 AT

twenty miles east of Durango. These specimens are now part of the
Strecker Collection at the Chicago Natural History Museum. The male
was apparently misidentified at one point in time and appears to be S.
cybele carpenteri (Edwards) (see Brown 1957a). It was originally re-
corded in the McCauley report (1879) as Argynnis cybele. The female
records were checked by L. P. Grey and one is figured in the report
(Plate I).

The flight span of S. n. nokomis is generally during the first three
weeks in August, although it has been taken from mid-July (Unaweep
Canyon ) into September.

Speyeria nokomis nitocris (Edwards )

Original Description: “Description of new species of diurnal Lepidoptera found in
North America.” Trans. Amer. Ent. Soc. 5: 15-16 (1874). Figured by Edwards,
B. N. A. 3: [91-92], Argynnis I; Holland, B. B., p. 85, pl. 13. (type).

Type Locality: White Mountains, Arizona. Based upon the discussion in Brown
(1965), and the study by the senior author of the distribution of nokomis in
Arizona, the type locality is probably east of Ft. Apache, along the East Fork of
the White River, Apache Co., Arizona. The lectotype (designated by dos Passos
and Grey 1947) (CM) bears the label “Nitocris ¢ W. Mtns. Ariz. type” and is
the specimen figured by Holland.

Discussion: The accompanying plate clearly shows the differences be-
tween n. nokomis and n. nitocris. Generally the colors are much more
intense in n. nitocris. In the males, ventrally, the discal area of the sec-
ondary is a deep cinnamon brown, the silver spots are quite brilliant and
heavily margined with black scales. The submarginal band is darker than
in n. nokomis and shows an orange flush over the lighter buff ground
color. In the females, dorsally, the dark colored areas are more extensive
than in n. nokomis and the blue-green overscaling is more prominent.
Ventrally, the discal area of the secondary has a cinnamon ground color
which is heavily overscaled with velvety black. The submarginal band
is yellow-buff but overscaled with dark scales.

The habitat of S. n. nitocris lies in lush Canadian Zone meadows or
along the mountain streams which feed such meadows. The adults show
a preference for feeding upon red thistles of various species. S. nokomis
nitocris generally flies at higher elevations that does S. n. nokomis. It has
been taken from 5400’ to 8500’ and appears to be more common at higher
elevations. It has been recorded from Arizona: Gila, Coconino, Green-
lee, Navajo, Apache Cos.; from New Mexico: Catron, Valencia, San
Miguel, Taos Cos. S. n. nitocris is found in the region of the Mogollon
Rim and White Mountain country in Arizona, and in the extension of this
terrain into New Mexico (Mogollon Mountains and eastward). It then

48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

COLORADO

FLAGSTAFF

O4

PHOENIX Ter + wovoe

ARIZONA
NEW MEXICO

Fig. 1. Probable distribution for S. nokomis. Solid dots indicate S. nokomis
nitocris; open dots indicate S. nokomis nokomis. Not all collection sites are shown.
The dotted line which encloses each series of dots indicates the probable range of
the given subspecies where suitable habitats exist. Mt. Sneffels is represented by the
open triangle and Ojo Verde by the solid triangle.

occurs in pockets at various other locations in New Mexico. The flight
period extends from mid-August into early September.

Of the two subspecies, nitocris is the more common. Both are quite
local, but nitocris has been taken in numerous localities and is some-
times found in high concentration. In both subspecies the males appear
to outnumber the females by a ratio of more than 5:1. The females ap-
pear less active than the males and keep to areas with dense vegetation.
In nitocris, the males appear as soon as the dew starts to dry off in the

VOLUME 25, NUMBER 1] 49

morning and they may be seen feeding or patrolling their territories. The
females appear more toward noon and except when feeding, are gen-
erally flushed from the tall grass and willows which border the streams
associated with their habitat. Mating behavior has been described by
Ferris (1969).

It is interesting to note that as one examines western populations of S.
nokomis through Utah and into California, the ratio of the sexes becomes
nearly equal.

From a somewhat limited study, it would appear that both subspecies
appear in strength in any given colony on an alternate year basis.

For comparison, also illustrated in the plate are specimens of S. nokomis
apacheana (Skinner) from California. Ventrally these are quite distinct
from the Rocky Mountain Region-Northern Arizona-New Mexico ma-
terial. In the males the discal area is almost the same color as the sub-
marginal band. In the females, the discal area is concolorous with the
submarginal band, but is overscaled with greenish-black.

We interpret the range of S. nokomis nokomis as being restricted to
southwestern Colorado, and to Uintah and San Juan Cos., Utah. It would
appear from preliminary studies that n. nokomis and n. apacheana inter-
grade in Utah. An interesting note on the blend zone situation was pre-
sented by Swisher and Morrison (1969). They collected nokomis from
California to Colorado. Based upon a somewhat limited number of
specimens, they found n. apacheana to be the only form in California.
In Washington Co., Utah (SW Utah at Leeds), they found 24% inter-
mediate between n. apacheana and n. nokomis. Collecting at Glendale,
Kane Co. (40 miles east of Leeds) yielded 17% intermediates and 15%
referable to n. nokomis. In northeastern Utah at Vernal, Uintah Co., they
found 20% intermediate and 44% were referable to nokomis. Colorado
specimens (Montrose Co.) were all n. nokomis. The remaining percent-
age figures at each site apply to n. apacheana. The sex ratio, males to fe-
males, are reported as 3:2 for n. apacheana and 9:1 for n. nokomis as an
average over the sites visited. Females of n. nokomis are quite scarce in
Colorado.

The range of S. nokomis nitocris is restricted to northern Arizona, New
Mexico and south-central Colorado along the New Mexico border. We
feel that S. nokomis nigrocaerulea (Cockerell and Cockerell) should be
treated as a synonym of S. nokomis nitocris because it evidently represents
an eastern outpost of the latter subspecies. Occasional New Mexico
specimens are larger than material from the White Mountains of Ari-
zona, and sometimes more darkly marked, but we do not feel this varia-

50 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Wi Aa
V-0% Wy
4 yy

Fig. 2. a-d. S. nokomis nokomis, Unaweep Canyon, Mesa Co., Colo., 6000’.
b, c, Female, 20-viii-1967; c, d, male, 18-viii-1967.

e-h. S. nokomis nitocris. e, Female, Willow Creek, Catron Co., New Mex., 25-
viii-67, 8100’; f, female, Pine Creek, Gila Co., Ariz., 7-viii-67, 5400’; g, male, Al-
pine, Apache Co., Ariz., 17-viii-67, 8300’; h, male, Pine Creek, Gila Co., Ariz., 7-
vili-67, 5400’.

i-l. S. nokomis apacheana, Round Valley, Inyo Co., Calif., 5000’. i, j, Female,
24—viii- 1968; k, 1, male, 11-viii-1963.

VOLUME 25, NUMBER l Dil

tion sufficient to warrant nomenclatural distinction. The New Mexico
insect was originally described from Beulah. This has created some
confusion as Beulah does not currently exist as a town. It is a mail route
in the Sapello Valley, San Miguel Co., New Mexico. Farming and ranch-
ing have essentially wiped out the habitat for nokomis. We would in-
clude here the specimen taken by Rotger in Conejos Ce.. Colorado and
described in Brown (1957b).

In some species of Speyeria, there is considerable change in discal color
intensity as a function of altitude. S. mormonia is a good example. We
considered this to be the situation with S. nokomis. Examination of col-
lection sites indicates that both n. nokomis and n. nitocris are found at
comparable altitudes although the latter extends into higher altitudes.
The ecologies of the respective habitats are quite different. It is on
basis of fascia, habitat, and geographical location that we have separated
the two subspecies. It should be noted that the underside color of n.
nitocris fades quite rapidly on the wing, and this may have been the
reason for confusing the two subspecies in the past. The very dark color
of the disc in fresh n. nitocris is most distinct from the color of the same
area in fresh n. nokomis.

We would thus propose a revised treatment of S. nokomis as follows:

SPEYERIA (SPEYERIA ) Scudder, 1872

Type: Papilio Nymphalis Phaleratus idalia Drury, “1770” [1773]
Argynnis Auctorum
=Dryas Hubner, [1806] (Opinion 278, name 75) (partim)
Neoacidalia Reuss, 1926 ( partim)
612 nokomis (Edwards ), “1861-3” (1862)
an. nokomis (Edwards), “1861-3” (1862 )
form valesinoidesalba (Reuss), 1926 (emendatio )
b n. nitocris (Edwards ), “1874-6” (1874)
nigrocerulea ( Cockerell and Cockerell), 1900
ab. rufescens (Cockerell ), 1909
nitrocris dos Passos, 1964 (lapsus calami )
cn. coerulescens ( Holland), 1900:
cerulescens McDunnough, 1938 (lapsus calami)
dn. apacheana (Skinner), 1918
nokomis (Edwards) 1873 (nec Edwards, “1861-3” [1862] )
e n. wenona dos Passos and Grey, 1945

Acknowledgments

The authors would like to thank the following collectors who supplied
specimens for examination, and information on habitats and ecology: J.
D. Eff, Bruce Harris, J. H. Masters, and R. E. Stanford. Scott Ellis pro-
vided extensive habitat notes for Colorado. Special thanks are due F.
Martin Brown of Colorado Springs, Colorado and L. Paul Grey of Lin-

JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Ol
bo

coln, Maine who provided collection data, reviewed the manuscript, and
made many pertinent comments during the preparation of this paper.
Mr. W. H. Shaughnessy of the Academy of Natural Sciences, Philadelphia
kindly provided data for specimens collected by Skinner in New Mexico.

Mr. Kilian Roever of Phoenix, Arizona kindly provided additional Ari-
zona collection records and made some additional comments upon the
habits of n. nitocris. He noted that females of this species generally ovi-
posit in shaded or semi-shaded open woods where there are seeps, al-
though one female was observed ovipositing in an open meadow formed
by an old beaver dam. Also supplied were records of specimens refer-
able to n. apacheana from Garfield, Iron, Kane, and Washington Cos.,
Utah. The intermediate situation between n. nokomis and n. apacheana
was noted as discussed above.

Literature Cited

Brown, F. M. 1957a. The McCauley expedition to the San Juan region of Colo-
rado in 1877. J. N.Y. Ent. Soc. 65: 139-145.

1957b. Colorado Butterflies. Denver.

1965. The types of the Nymphalid butterflies described by William Henry
Edwards Part I. Argynninae. Trans. Amer. Ent. Soc. 91: 233-350.

pos Passos, C. F. 1964. <A synonymic list of the Nearctic Rhopalocera. Lep. Soc.
Mem. I.

pos Passos, C. F. AND L.. P. Grey. 1947. Systematic catalogue of Speyeria (Lepi-
doptera, Nymphalidae) with designations of types and fixations of type locali-
ties. Novitates 1370.

Epwarps, W. H. 1868-1872. The Butterflies of North America. Volume 1.
American Ent. Soc. Reprinted 1888 by Houghton Mifflin and Company, Bos-
ton.

1874-1884. The Butterflies of North America. Volume 2. Houghton

Mifflin and Company.

1887-1897. The Butterflies of North America. Volume 3. Houghton
Mifflin and Company.

lerris, C. D. 1969. Some additional notes on mating behavior in butterflies. J.
Lepid. Soe, 23:3) 27272,

HoitaAnp, W. J. 1898. The Butterfly Book. Doubleday, Page and Co., New York.

Macoms, J. N. anv J. S. Newserry. 1876. Report of the exploring expedition
from Santa Fe, New Mexico, to the junction of the Grand and Green rivers of
the great Colorado of the West, in 1859. Washington.

McCautey, C. A. H. 1879. Report on the San Juan Reconnaissance of 1877, by
Lieutenant C. A. H. M’Cauley, Third Artillery, in charge. In Index to the
xecutive Documents of the House of Representatives for the Third Session of
the Forty-fifth Congress, 1878~79. In 18 volumes. Volume V. Report of the
Chief of Engineers, Part III. Washington.

SwisHeR, W. L. AND A. L. Morrison. 1969. News of the Lepid. Soc., 15 April,
p. 4.

VOLUME 25, NuMBER 1 53

OBSERVATIONS ON THE BIOLOGY OF OCNEROGYIA
AMANDA STGR. (LYMANTRIIDAE), A PEST OF FICUS
IN IRAQ

PETR STARY
Institute of Entomology, Czech Academy of Science, Praha

IBRAHIM K. Kappou

Biological Research Centre, Council of Scientific Research, Baghdad

The comparison of the present observations with data in the literature
has shown that there is a considerable lack of information about the moth
pest, Ocnerogyia amanda Staudinger on figs in Iraq. It is intended, in
this paper, to cover at least some of these existing gaps.

This work was carried out in Baghdad from April to July, 1968, both in
the field and in the laboratory.

The eggs are laid in groups of about 20 or more. Similar data was ob-
tained by Buxton (1920) and Ramachandra (1922). Most eggs can be
found on the lower side of the leaves and to a lesser degree on the upper-
side of leaves, on the trunk or on dry leaves under the trees.

The newly-hatched first-instar larvae disperse over the leaf on which
the eggs were laid and begin feeding. Movement of larvae is limited
while small but becomes more extensive as they grow older. The feeding
behaviour of the larvae varies with their size. First-instar larvae skeleton-
ize the leaf surface; second-instar larvae make small holes in the leaves;
older larvae eat great holes in the leaves or devour them completely ex-
cept for the main ribs. The early-instar larvae are not noticeable on the
tree, but those of later instars are conspicuous because of their greater
size and long yellowish body hairs.

The larvae exhibit a definite diumal migration. They feed at night
but take shelter among dead leaves or soil beneath the fig trees during
the day (Buxton, 1920; Ramachandra, 1922). According to Scott (1929),
fig trees neighbouring mud walls were severely attacked because larvae
thrive where they can find shade at certain times of the day; moreover
larvae feed on the lower leaves during some part of the day whereas
they feed on the more exposed foliage at night. It was found during this
study that the larvae do not feed during the hot periods of the day. Dur-
ing this period, they congregate in places which are cooler and more
sheltered than are the leaves. Such places are parts of the trunk, espe-
cially near the base, and sheltered parts of garden walls. Both young
and older instar larvae may be found in such aggregations. The larvae

?
JOURNAL OF THE LEPIDOPTERISTS SOCIETY

iggs of Ocnerogyia amanda Ster.

Damage to fig leaves by larvae. 2, By first-instar larvae; 3, by second-
yy late-instar larvae

VoLUME 25, NuMBER 1

instar larva; 6, cocoon.

Fig. 7. Congregation of larvae at the base of a trunk

56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

were observed to feed in the late afternoon during July. It is possible,
however, that during the hottest periods of the year feeding is restricted
to the hours of darkness, especially on more exposed and unprotected fig
trees.

According to Ramachandra (1922), the larvae have six moults. Larval
and pupal stages last about 24 days and 8-11 days respectively. Scott
(1929) reported somewhat similar results, the egg, larval and pupal
stages being 5-6, 21 days and about a week respectively.

The cocoon is spun of rough yellowish-grey silk. The texture is irreg-
ular and the pupa is fully visible in its cocoon. Laboratory observations
have shown that the larvae do not spin the cocoons on plain surfaces, but
only on the sides or on the top of the cage. In nature, they do not occur
on green leaves, but can be found on shaded trunks, usually the lower
parts, and in dry leaves. Other preferred sites are sheltered parts of gar-
den walls. Usually, congregations of pupae occur in suitable places. The
congregation of the pupae is similar to that of the larvae; the old larvae
select obviously microclimatically favourable sites for pupation, whereas
the younger larvae occur in such places only temporarily and migrate to
their feeding sites. The number of pupae observed in one congregation
is usually three to five. The distance from a tree is usually not more than
about five meters.

The adults can be caught easily in light traps in late evening hours.
There is sexual dimorphism, the female being light brown while the male
is somewhat smaller in size and with orange hind wings.

The seasonal occurrence of the moth was noted to be continuous
throughout the summer. This is considered as unusual among Mesopo-
tamian Lepidoptera, which generally aestivate during summer (Buxton,
1920). It was found during the present investigation that adults occur
in the last 10 days of May. The eggs and young larvae were observed
during the first half of June and the pupae and adults at the end of June
and in the first half of July. Obviously, there are at least two generations
and perhaps three a year. Ramachandra (1922) similarly reported the
duration of the entire life cycle as being about one and one half months
and that there are probably three generations a year.

The lack of aestivation in this moth during the unfavourable hot sum-
mer months is obviously permitted by the ability of the larvae to migrate
to protected shelters and return to feeding sites at times when conditions
are cooler.

O. amanda is very important economically causing serious damage to
tig trees. Larvae feed on the leaves and sometimes devour them com-

VoLUME 25, NuMBER 1 51

pletely. Often the greater parts of trees were observed to be practically
leafless with only the ribs left and none of the leaf tissues. The shriv-
ellng and dropping of fruits may be also observed, and is caused by a
sequence of leaf damage and water-regime to the damaged tree. Other
factors may also be responsible for it, such as the exposure of fruits to
direct hot sunlight and, in some cases, infestation by a fly, which stunts
the growth of fruits and causes their shedding.

According to Buxton (1920), this pest should be fairly easy to keep in
check; the Arab growers burn the dead leaves and other rubbish in heaps
beneath the trees. This destroys all stages of the insect as well as its hid-
ing places. But this study has shown that such a control method is not
sufficient because burning does not destroy all individuals of the pest; the
trunks of the trees and garden walls are not touched. More recent meth-
ods should provide adequate control of this insect pest. If the population
of larvae reaches economic levels, treatment by insecticides must be
undertaken during that period of the day when the larvae are present
and feeding on the trees—i.e. late afternoon, evening and night.

Acknowledgments

Sincere thanks are due to Dr. J. Moucha of the National Museum at
Prague for identification of the insect species and for further valuable
information. The writers wish also to express their gratitude to the Bio-
logical Research Centre of the Iraqi Council of Research for providing
the space, facilities and the opportunity to carry out this work.

Literature Cited

Buxton, P. A. 1920. A Liparid moth, Ocnerogyia amanda Staud., destructive to
figs in Mesopotamia. Bull. ent. Res. 11: 181-186.

RAMACHANDRA Rao, R. S. Y. 1921. <A preliminary list of insect pests of Iraq.
Dept. Agric. Iraq Memoir 1: 28-29.

RAMACHANDRA, Rao (Y.). 1922. Notes on the life-histories of two Mesopotamian
moths. Bull. ent. Res. 12: 477-479.

Scotr, H. 1929. Notes on the life-history of the fig-tree moth, Ocnerogyia amanda
Staud. (Lymantriidae). Bull. ent. Res. 20: 39-40.

Wimsuurst, C. R. 1920. Entomological Section. Administration Report Agric.
Directorate (Mesopotamia) for 1919, pp. 39-41. Baghdad.

5 OURNAL OF THE LEPIDOPTERISTS SOCIETY
2)

THE LIFE HISTORY OF ORNITHOPTERA ALEXANDRAE
ROTHSCHILD

R. STRAATMAN

Department of Agriculture, Stock and Fisheries, Konedobu, Papua/New Guinea

During the years 1967 and 1968 some data were obtained on the life
history and behaviour of Ornithoptera alexandrae Rothschild, the largest
known species of Rhopalocera; the study was completed in 1970. The
geographic distribution of O. alexandrae is limited to a relatively small
area in southeastern New Guinea. However, within its range there are
many areas where the butterfly does not occur although the hostplant
grows prolifically. The main habitat is a low and relatively flat region,
but it has also been observed at altitudes up to 900 meters.

O. alexandrae is monophagous. Its hostplant is Aristolochia schlechteri,
a vine having rather large, thick leaves and stringy stems covered with a
layer of strongly ribbed cork. The flower is shaped like a starfish with
three long arms and is dark purple-brown with a yellow heart. The fruit
is green, shaped like a small cucumber, 20 to 30 cm long, strongly ribbed
longitudinally and has a rough skin. It matures slowly and when fully
rotten the seeds fall to the ground and are carried away by rainwater gen-
erally over short distances, resulting in a number of plants growing in a
restricted area. In primary forest the vine reaches the top of tall trees of
over 40 meters high. When larvae were transferred to Aristolochia tagala,
a plant more generally distributed, it was accepted readily and the larvae
developed normally, although at a much faster rate than larvae feeding
on. their natural host. Data recorded show a rapid growth as was the case
with larvae of Papilio aegeus when reared on parsley or carrot leaves
(Umbelliferae) instead of their natural foodplants (Rutaceae) (Stride
& Straatman, 1962). When three larvae, obtained from eggs collected in
the field, were reared on A. tagala they went through six instars instead
of the usual five. It is not known whether this is hereditary or environ-
mental. The female butterfly does not oviposit on A. tagala.

Oviposition. Generally a single egg is laid on the under surface of an
old leaf of the hostplant. In secondary forest where this plant is not very
tall, the egg is laid from a few centimeters above the ground to about
one meter above it. On several occasions a female was observed laying
on other objects than the foodplant such as a grass stem growing at a dis-
tance of a few centimeters from one of the main stems of the Aristolochia

ine. In primary forest, however, oviposition may take place at a con-

le height above the ground.

VoLUME 25, NuMBER 1 59

Egg. Large, light yellow, flattened at the base. Diameter 3% mm. Covered with
a thick layer of a bright-orange sticky substance, which fixes it firmly to the surface
on which it is laid. Incubation period varying from 11 to 13 days.

First-instar Larva. Ground colour dark wine red. All segments with long tubercles
of same colour as body; tubercles fleshy for about one-fourth their length, remain-
ing part stiff and black with numerous black spines. Two dorsal tubercles on the
fourth abdominal segment light red as is dorsal saddle mark joining them on the
same segment. Saddle mark divided mid-dorsally by a narrow black line. Head,
prothoracic shield and legs black; prolegs dark, fleshy. Newly hatched larva seven
to eight mm long. Osmaterium orange yellow.

Second-instar Larva. Ground colour reddish black. Tubercles proportionately
longer, all fleshy and without spines, latero-dorsal ones the longest. Dorsal and
latero-dorsal tubercles on thoracic segments two and three, and dorsal ones on ab-
dominal segments one, seven, eight and nine red; two dorsal tubercles on ab-
dominal segment four creamy-white with pink tips; remaining tubercles of ground
colour. First thoracic segment with four tubercles, following three segments with
eight; abdominal segments two to eight with six; ninth abdominal with four and the
last segment with two tubercles.

Third- to final-instar Larvae. Ground colour unchanged. Tubercles without spines,
of nearly equal length except for the ventro-lateral ones which are very short. In
ultimate instar, body tubercles proportionately smaller than in early instars. All
bright red except two dorsal ones on fourth abdominal segment which remain creamy
white with pink tips. Conspicuous saddle mark extending and narrowing down to
spiracles. Some larvae with an additional creamy spot on third abdominal segment.
Measurements of a large, mature larva: length 118 mm, greatest width 30 mm;
headcapsule length, 12 mm, width 11 mm; longest tubercle, 13 mm. Some larvae have
six instars instead of the usual five, and these producing the largest butterflies.

Adult. The size of the butterfly varies considerably. The average length
of the forewing in the male is 97 to 100 mm, in the female 115 to 126 mm.
Many specimens are smaller, few are larger. One previously unrecorded
feature is that some males have translucent, yellow discal spots on the
hind wings, homologous to those of priamus and victoriae.

Duration of stages. The following data were recorded for the average
duration of each stage:

on Aristolochia schlechteri on Aristolochia tagala
egg 12 days egg 12 days
first instar 5 days first instar 3 days
second instar 7 days second instar 6 days
third instar 9 days third instar S days
fourth instar 11 days fourth instar 9 days
fifth instar 20 days fifth instar 10 days
sixth instar 22 days sixth instar 14 days
prepupa 3 days prepupa 3 days
pupa 42 days pupa 42 days

Total 131 days Total 107 days

60 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Ornithoptera alexandrae Roths., first-instar larva.
Fig. 2. O. goliath Oberth., first-instar larva.

/

Figs. 3, 4. O. alexandrae. 3, Second-instar larva; 4, third-instar larva.

VoLUME 25, NuMBER 1] 61

Figs. 5-7. Ornithoptera alexandrae Roths. 5, Sixth-instar larva; 6, pupa, lateral;
7, pupa, dorsal.
Fig. 8. O. victoriae epiphanes Schmid, pupa, lateral.

62 JouURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 9. Pupae of Ornithoptera alexandrae Roths. (below) and O. priamus poseidon
Dbhdy. (above).

Figs. 10-12. O. alexandrae, adults. 10, 12, Male; 11, female.

In the higher altitudes of its range the total figure may reach 180 days
depending on the locality.

It was observed that larvae feeding on Aristolochia schlechteri spend
much time in search of suitable leaves and stems. However, those feed-
ing on the succulent and soft parts of Aristolochia tagala rarely move
around and appear to have a longer average daily feeding time.

Feeding habits. Shortly after hatching the larva devours its eggshell,
which provides sufficient food for the next 24 hours. It then commences
feeding on tender shoots and young leaves. Later instars feed on older
leaves and stems. Fifth and sixth instar larvae feed mainly on the stringy
stems and shortly before pupation one or more stems of the host vine
are severed, causing the upper parts to wither. If the plant is young, the
lower part is eaten down to the ground.

Pupation. The larva may wander for 24 hours or longer to locate a
suitable site for pupation, which sometimes occurs at a considerable dis-
tance from where it was feeding last. The longest recorded distance was
nearly 10 meters. It generally pupates under a leaf of any kind of shrub
or tree other than the hostplant, rarely on stems, at an average height of

VoLUME 25, NuMBER 1 63

one to two meters above the ground in secondary forest, but considerably
higher in primary forest.

Pupa. Ground colour light brown. Wing cases yellow, a broad light-brown streak
along lower margin. Abdominal segments brown ventrally and yellow dorsally, with
a yellow latero-ventral streak. Dorsal saddle mark bright yellow, extending over
segments one to five. Middorsally divided by a narrow dark brown line; a similar line
running laterally below wing cases. Thorax dark brown, teguiae bright yellow.
Abdominal segments five to eight with two very short, sharp, black processes each.
Pupa very closely resembling that of O. victoriae. Duration of pupal stage from
forty to forty five days.

General observations. Female butterflies appear to follow a deter-
mined flight pattern when ovipositing. This is suggested by the fact that
larvae in different stages of development, together with one or more
pupae or exuviae, may always be located on or near the same foodplant,
while other plants growing in the vicinity remain free of specimens the
year round. It is possible that females, in their search for suitable con-
ditions for oviposition, are stimulated by plants which already do sup-
port or have previously supported early stages. Single larvae are only
found on small plants.

Although it has not been possible to determine the longevity of the
adults it is believed to be similar to that of O. priamus. Some males of
alexandrae were clearly marked after they emerged from the pupa, and
then released in a garden. A few of these specimens established them-
selves for the duration of their adult life in the same garden, where both
hostplants and flowers were prolific. After 11 weeks, one male was
found caught in a large spiderweb. Another died in the same way after
nearly 12 weeks.

Predators and diseases. Several larvae have been found marked with
numerous rustbrown dots, sitting motionless without feeding. They died
after several weeks and may have been killed by a fungus disease.

On one occasion a larva covered with eggs of a Tachinid was collected,
but continued feeding, pupated normally and produced the adult. Ap-
parently the parasite eggs did not hatch or the resulting maggots did
not survive. A native collector once had a pupa which produced a large
number of small, black wasps, probably Chalcididae. Otherwise larvae
of O. alexandrae are rarely attacked by parasites. Prepupae and soft.
fresh pupae are sometimes killed by ants and wasps and mature larvae
and pupae are attacked by tree rats and small marsupials. When not
mating both sexes may be seen, generally flying high (average 20 to 30
meters above the ground) and in a single direction. In the forest, males
are seldom seen as they remain in the shade of the high canopy and avoid
open or exposed areas.

64. JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fecundity. Two females were taken on the wing while ovipositing.
Both specimens appeared in good condition, but it was impossible to
make an accurate estimate of their age. They were kept alive and fed
daily with a sucrose and honey solution. After 12 days in captivity both
specimens were killed. One female had laid one egg; dissection of its
abdomen produced another 16 mature eggs. On the basis that not more
than 10 eggs had been laid before its capture, we can estimate a total
capacity of 25 to 27 eggs. Dissection of the second female produced a
total of 12 eggs of which two were immature.

Acknowledgments

Thanks are due to Dr. J. L. Gressitt, Bishop Museum, Honolulu for
his assistance in the publication of the photographs and to Mr. Ted Fen-
ner, Department of Agriculture, Stock and Fisheries, Konedobu, Papua,
for critically correcting and improving this manuscript. The work was
partly supported by National Science Foundation grant GB-7330 to the
Bishop Museum, Honolulu. Dr. F. Schmid, Entomology Res. Institute,
Ottawa, prepared the plates, and corrected the manuscript and galley
proofs.

Literature Cited

Jorpan, K. 1908. Indo-australian butterflies. In Seitz, A. Macrolepidoptera of
the World. 9: 12-13.

STRIDE, G. O. AND R. STRAATMAN. 1962. The host plant relationship of an Aus-
tralian swallowtail, Papilio aegeus, and its significance in the evolution of host
plant selection. Proc. Linn. Soc. N. S. W. 87: 69-78.

A HOST PLANT FOR NORTHERN POPULATIONS OF EUCHLOE OLYMPIA
(PIERIDAE )

Following my report in the Annual Summary for 1967 of Euchloe olympia (Ed-

wards ) being sighted in large numbers between Deep River and Stonecliff, some
75 miles farther north and possibly east of the Killaloe, Ontario records reported in
the 1966 Annual Summary, I unsuccessfully attempted during the succeeding years
to discover the host plant of this northern population.
_ It was not until May 23 of this year, that I was fortunate enough to observe two
females of Euchloe olympia, at 1430 hours and a temperature of 76° F., ovipositing
on Arabis glabra (1..) Bernh. This plant is regionally (although probably incorrectly )
called Tower Mustard. Doubtless other Cruciferae are fed upon by olympia in this
area but these are yet to be determined.

Identification of the host plant was very graciously made by Mrs. Mary Moore of
the Petawawa Forestry Station, Canada Department of Fisheries and Forestry.

5. A. Linpsay, Deep River, Ontario.

VOLUME 25, NUMBER 1 65

PAPERING LEPIDOPTERA IN GLASSINE ENVELOPES

RIcHARD E.. GRAY
RFD No. 1, Enfield, N. H.

In several previous volumes of the Journal, especially in the late fifties
and early sixties, there appeared articles on papering Lepidoptera. Many
of these recommended using paper triangles instead of glassines because
of certain defects in usage of glassines. It is my purpose in this short note
to describe my usage of glassine envelopes and to give a few tips on how
to avoid some of the difficulties mentioned by others.

I use glassine envelopes exclusively in papering all Lepidoptera cap-
tured by me, for the following reasons: ease of use and visibility.

Ease of use. I find it difficult, when trying to use triangles, to get the
flaps to stay down. I also dislike to have to prepare triangles. I have
also found that in certain small, fat bodied skippers and in many sphingid
and arctiid moths the trouble getting them to stay in place while folding
the flaps over can cause the specimens to become rubbed or to slide out of
place. With glassines there are no such problems. The glassines I use
have only one flap (along the top edge), and with my usual size glassines
(see below), I have only to fold the flap over and make a good stiff
crease. The flap stays folded. Also, the pressure that is applied to the
specimen by the front and back of the envelope, sealed as it is on three
sides, holds the specimen in place without slippage.

Visibility. This is the most important quality the glassine envelope has
over the triangle for most collectors. It allows you to see exactly how
the specimen is placed in the envelope and also allows you to determine
the condition of the specimen without having to handle it more than
necessary.

How to select the proper glassine to use. First, use only glassines with-
out gummed flaps. Many collectors prefer to use glassines with gummed
flaps because they can be ticked shut. Unfortunately, they can also be
“licked shut” by humidity in the atmosphere. I have found that if after
papering a specimen in a glassine that has no gum on the flap, I fold
the flap over, lay it on a piece of pasteboard or other firm, flat surface,
and run a curved, smooth surface over the crease of the flap that the
resulting stiff crease will not unfold. Secondly, use the smallest size of
glassine that the specimen will fit into without touching either of the
two sides and without coming into contact with the crease of the flap
(remember that you will be applying pressure to the flap crease). For

66 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

most lycaenids, pierids, geometrids, and hesperids, as well as other Lepi-
doptera of this approximate size I prefer the standard two inch by two
inch glassine coin envelope. These come with the flap uncreased and
should cost four to five dollars per thousand at the most. For larger
specimens such as Danaus plexippus (Linnaeus) or Limenitis archippus
(Cramer), or for Papilio I use the various standard size postage stamp
envelopes. The sizes to use will depend on the size of the specimen. The
amount to buy will depend on which butterflies and moths you are col-
lecting, or which are commonest in your area and are collected most. I
suggest that you purchase a selection of sizes ranging from number ones
(one and three-quarters by two and seven-eighths inches), to number
fours (three and one-quarter by four and seven-eighths inches). If you
need larger sizes they are available. Try out the different sizes and see
which you use most. Once you know which size or sizes you use the most,
buy these by the thousand. They are much cheaper this way.

How to use your glassines. This will vary with individual preference.
I use my glassines as follows. When I go out collecting, I carry in an
upper shirt pocket about one hundred number four glassines. In front
and in back of this pile of glassines are pasteboard separators from the
boxes of one thousand that I have bought. The separator closest to me
prevents perspiration from coming in contact with the glassines. This
is important since glassines will wrinkle when wet. The front separator,
the one furthest away from me, is used to divide the empty glassines from
those containing specimens. Having caught a specimen in my net, I
“pinch” it and then remove it from the net with my forceps. Holding
the forceps in one hand, I reach into my pocket and between the two sep-
arators with the other and remove an empty glassine, opening it one-
handed as I remove it; this takes practice, but not much. I then insert
the specimen into the envelope with the forceps, put away the forceps,
spread the separator and the pocket front apart with two fingers of one
hand and carefully insert the glassine containing the specimen with the
other hand. Once you have practiced this technique a few times it will
become almost automatic. Take care, however, never to “cram full” the
pocket. If it becomes a tight fit and a glassine does not go in smoothly
simply transfer some of the full glassines elsewhere.

Having collected enough specimens to make a good day, I return home
and, removing the glassines, place them in piles according to species. I
then prepare the permanent storage glassines.

Preparing permanent storage glassines. Having removed the speci-
mens I wish to discard or spread, I then separate each pile, one pile at

VoLUME 25, NUMBER 1 67

a time, by sex. I now take my two by twos and put the appropriate data
on the back flap of each envelope. I use a nylon tipped pen with a quick,
surface-drying ink. You may, if you wish, use a typewriter or rubber
stamp, but since glassines are not porous you will have to wait some
time for the ink to dry. With a surface-drying ink such as is used in felt-
tipped marking pens you eliminate the waiting. By the time you finish
the last envelope in the species from one pile, the first will be well dried.

Putting the specimen in the glassine. Using my forceps I remove the
specimen from the number four envelope and place it in the prepared two
by two with the antennae in the usual “down between the wings” posi-
tion. Carefully removing the forceps I adjust the flap to the proper
height and put a light crease on the edge with the forceps. I then, as
described above, put a permanent stiff crease on the edge. The specimen
is then entered into the record book and placed in permanent or tem-
porary storage.

Storing specimens. Many people use cigar boxes or other containers
for storing papered Lepidoptera. With my system of two by twos how-
ever, I can use plastic coin storage boxes with tight fitting lids. These
boxes, approximately two and one-quarter by two and one-quarter by
nine inches are made specifically to hold two by two inch coin envelopes.
It is possible to arrange the specimens in any order I wish, add an in-
secticide, put the top on and tape the box shut, and put the sealed box
away in storage for years without the necessity of further care. Labels
are available to go on the ends of the storage boxes, thus the contents are
recorded and the proper box can be retrieved with little effort.

Warnings. Never use an insecticide in a plastic box that will react
chemically with the plastic. Paradichlorobenzene and other oil based
insect killers are verboten! I use a powdered insecticide and fungicide
combination such as is used in many museums. This works just fine.
Another precaution is to allow all specimens two to three weeks to dry
out thoroughly before sealing the box and to make sure that as much
moisture as possible is excluded from the box. I suggest sealing boxes on
the second of two dry days or in winter when the moisture content of
the air is low. In the tropic and humid regions I recommend the use
of warm air to dry out the box before filling. Use your own judgment
on how to go about it, but never heat the box in a drying oven. At best
it will probably warp and at worst it will melt.

Relaxing specimens in glassine envelopes. As stated above, glassine
envelopes are non-porous. They cannot, therefore, be just placed in a
relaxing box like triangles can. A simple method of relaxing such glas-

68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

sined specimens is as follows. Take the envelope, and with a pair of good
straight scissors cut off the smallest bit you can manage all along both
sides and also along the top. Cut as close to the specimen as possible,
leaving extra glassine above the specimen. Taking the extra glassine in
the forceps, thus keeping the specimen immovable between the remain-
ing part of the glassines back and front, place the specimen and remains
of the envelope in the relaxing box. The glassine will curl from the
humidity and expose the specimen on three sides to the atmosphere in
the box. The specimen will not be “soaked” if the glassine is placed
directly on the wet sand or sponge you use to hold the moisture since
water will not penetrate the non-porous glassine.

If you have used a waterproof ink, and have put the data on the front
of the envelope you can make sure that the data is included on the glas-
sine that goes into the box with the specimen. If the data is on the flap
(which you cut off along the top to open the envelope), you may put
the flap in with the specimen or put it under the relaxing box, as I do.
Choose the method that best suits you.

When the specimen has relaxed, remove it using the extra glassine
again, and go to work.

The above are the techniques I use to get excellent results with glas-
sine envelopes, at the lowest expenditure of time, effort, and cash. Please
remember that circumstances vary and that modifications or substitutes
should be used to suit the individual collector.

HYBRIDS AMONG SPECIES OF HYALOPHORA

Davin ALLAN WRIGHT
18 Clinton Place, Woodcliff Lake, New Jersey

Although hybrids between species of Hyalophora are well known and
the triple hybrid (H. gloveri 6 x H. rubra 2) 6 X H. cecropia 2? has
been described (Collins and Weast, 1961), there does not appear to be
any description of the triple hybrid (H. cecropia 6 x H. gloveri 2) 6 x H.
rubra 2 or of a hybrid which combines all four species. The object of
this investigation was to raise the quadruple hybrid [(H. cecropia 6 x
1. gloveri 2°) 6 X H. rubra 2]8 X H. columbia 2 and to study the effects
of foodplants on the rate of growth and the size of cocoons and adults.

VoLUME 25, NuMBER 1 69

TABLE 1. Hyalophora hybrids. Comparative growth of larvae on different food
plants*

No. Surviving % Survival
——__—_————__ Larval Stage

Foodplant Number To Pupa To Adult To Pupa To Adult Mean Days
H. cecropia 6 X H. gloveri 2

Willow iG tS 14 88 82 42,

Cherry 25 18 16 12 64 54

Maple PAS: 20 18 80 72 US

Alder Sal 5 3 24 14 70
[(H. cecropia 6 x H. gloveri 9) 6 XH. rubra 9] 6 XH. columbia 9?

Willow (A) Deg D3 ie 85 48? AT

Willow (B) OT 12, 10” 44 Sie. Sy

Cherry 55 39 29° 71 Dom 44

Tamarack 20 D IL 10 5 48

2 Data on the trihybrid were incomplete and cannot be reported. However, survival was ex-
cellent, and the larval stage was about 35 days on both willow and cherry.

b Results are for those that had emerged within 12 months of pupation. Most of the others were
still alive and emerged the second summer.

Procedure

A male and a female were put in an eight cubic foot wire cage to secure
a mating. Copulation began in the early morning and continued until
the evening. The female was put in a paper bag and left about three
days to lay her ova. The ova were placed in a typewriter ribbon box and
hatched in about 10-15 days.

The larvae were fed washed leaves of the foodplant, the stems of
which were put through holes in the tops of plastic boxes containing
water. Most of the larvae were grown in fish tanks which were covered
with mosquito netting and kept indoors at a temperature which varied
from 20-30° C. The tanks were cleaned and the larvae were fed at fre-
quent intervals. In some cases larvae were grown outdoors in mosquito
netting sleeves or transferred to large wire cages in the fourth and fifth
instars to prevent crowding. The larvae either spun in their containers or
were put in paper bags to spin. Pupae were overwintered in a cool place.
Measurements were made of the size of the cocoons and the wingspans
of the adults. Characteristics of the larvae were recorded at each stage.

Discussion and Results
(see Tables 1 and 2)
H. cecropia é X H. gloveri ?

After the first instar, larvae were raised outdoors in sleeves. The larvae
grew fastest, had the lowest mortality, spun the largest cocoons, and

7 OURNAL OF THE LEPIDOPTERISTS SOCIETY
{

TABLE 2. Hyalophora hybrids. Size of cocoons and adults

Size of Cocoons (mm) Mean Wing Length (mm)#
Foodplant Mean Length Mean Circumference Males Females

H. cecropia 6 X< H. gloveri 9

Willow 54 88 62(7) 65(5)
Cherry 49 80 53(9) 59(7)
Maple 45 18 By 1) 55(5)
Alder 4] 67 44(1) Bye)
(H. cecropia 6 X< H. gloveri 2) x H. rubra @

Willow 65 100 60(16) 64(15)
Cherry 60 96 56(9) 60(9)
[(H. cecropia 6 <x H. gloveri 92) 6 x H. rubra 9] 6 < H. columbia 9?”
Willow (A) 58 80 55(4) 59(9)°¢
Willow (B) 49 70 50(7) BA (<3)
Cherry at 76 57(15) 60(14)
Tamarack 44 Gall — 53(1)

* Measured from the base to the apex of the right forewing. The number within parentheses
indicates the number of specimens measured.

» Wingspan of the female H. columbia was 50 mm, while that of the hybrid male was 59 mm.

© Results are for those which had emerged within 12 months of pupation.

formed the largest adults when raised on black willow (Salix nigra).
Black cherry (Prunus serotina) was the second most suitable foodplant,
sugar maple (Acer saccharum) was third and alder (Alnus) was poorest.

(H. cecropia 6 X H. gloveri 9) 6 x H. rubra @

Of 253 ova, 222 hatched from May 5 to 8, 1967. They were divided
into two groups. One was fed weeping willow (Salix babylonica) and
the other was fed black cherry. Because of the large number of larvae,
only a representative sample was carried to the adult stage. The larvae
on both foodplants spun in about 35 days. Both cocoons and adults were
slightly larger when the larvae were reared on willow, suggesting that
this is a slightly superior foodplant to cherry.

The adults (Figure 1) varied in color but had stronger H. rubra Boisdu-

val characteristics. The cocoons also strongly resembled those of the H.
rubra.

[(H. cecropia 6 x H. gloveri 2) 8 x H. rubra 2] ¢ X H. columbia °

A mating was obtained on May 13, 1968. The pair broke up at about
5:30 PM. A total of 129 ova were laid, all of which hatched from May 27
to 29. Twenty were put on tamarack (Larix americana ), 54 on weeping
willow, and 55 on black cherry. The tamarack was obtained from a peat

VOLUME 25, NuMBER 1 Gl

Fig. 1. Hyalophora hybrids. A, B, (H. cecropia 6 x H. gloveri 9) ¢
rubra 2, male and female; C, D, [(H. cecropia 6 «x H. gloveri 2) 6 « H.
2] 6 x H. columbia 2, male and female.

Seal.
rubra

bog in Sussex County, New Jersey. Two types of weeping willow were
used: one (A) was more limber and longer lasting, whereas the other
(B) was brittle and required more frequent replacement. These food-
plants were chosen because the hybrid and triple hybrid both did well
on weeping willow and black cherry. H. columbia Smith has been raised
on choke cherry and in nature feeds only on tamarack.

The adults (Figure 1) resembled giant H. columbia, but were more
reddish and somewhat lighter in color. Cocoon size and survival rate
were greatest for specimens grown on willow and cherry. The larvae
did not do well on tamarack and all but two died in the early instars.

Descriptions of the Larvae

(H. cecropia 6 < H. gloveri 2) 6 XH. rubra @

First instar: Bristly black. Appeared to be identical to other members of genus
Hyalophora.

Second instar: Ground color yellow. Double row of black spots between dorsal
tubercles (DT) on segments 2-10, upper lateral tubercles (ULT) and DT on seg-
ments 2-10, and lateral tubercles (LT) and ULT on segments 2-11. All tubercles
black with yellow base. Frons usually black but sometimes brown. Clypeus silvery
greenish, head otherwise black.

—l
bo

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Third instar: Ground color yellowish green but turquoise dorsally. Between DT
on segments 3-10, one black spot per segment, sometimes two on segment 10. Black
spots between ULT and LT on segments 3-11, ventral to spiracles. Sometimes a
black spot between ULT and DT on segment 2. Sometimes two per segment dor-
sal to each proleg and segment 10. Often no spots at all except one on each seg-
ment in line with lower lateral tubercles (LLT). Spiracles white or yellow, ringed
with black. Lobes yellow green with two black spots. Frons yellow green, clypeus
and labrum silver, palpi yellow green with one black setae, mandibles black. DT on
segments 2-10 and LT yellow. Other tubercles blue or black with blue base.

Fourth instar: Light to dark green, light turquoise dorsally. Segment 1 dorsally,
frons, labrum and palpi yellow green. Lobes yellow-green with black dots. Clypeus
blue, mandibles black. Spiracles white ringed with black. DT on segment 1 blue,
sometimes reduced to just a dot. DT on segments 2-4 yellow to orange with broken
ring of black dots. Other DT and LT yellow. Other tubercles all blue at base to
blue-white at tops.

Fifth instar: Green, lighter dorsally. Segment 2 dorsally, lobes and frons yellow
green. Frons somewhat yellower than lobes. Clypeus blue grey, labrum grey, palpi
turquoise, mandibles black. DT on segments 2-4 all had black ring (broken on
segment 4). DT yellow at base, orange or yellow above ring, and bulbous. Tuber-
cles on segment 1 rudimentary black bumps. DT on segments 5-10 and CT yellow.
CT slightly bulbous with broken black ring. Other tubercles blue at base to white
on top, with black ring around base.

[(H. cecropia 6 x H. gloveri 2) 6 x H. rubra 2]6 * H. columbia 2°

First instar: Same as other Hyalophora.

Second instar: Ground color yellow to greenish. Row of black spots dorsally one
or two per segment. Row of single spots between DT and ULT, and between ULT
and LT. Spiracles white with black ring. Head completely black.

Third instar: Mainly olive-green, but turquoise dorsally, except for yellow first
segment. One black spot on segments 3-9 dorsally. One spot per segment between
ULT and LT on segments 2-11 inclusive, two per segment between ULT and DT,
one spot per segment below LT on segments 4-12. All had lower two rows of spots,
but not all had upper two rows. Lobes green with three black spots, sometimes
fused together. Frons green, clypeus grey, labrum blue-grey, palpi yellow-green,
mandibles black. Spiracles yellow ringed with black, tubercles black. DT on seg-
ments 3-10 with yellow base. CT yellow anteriorly.

Fourth instar: Basically green, lighter green to blue dorsally. Lobes, frons, and
palpi yellow-green. Clypeus blue to grey, labrum grey to black, mandibles black.
Spiracles white ringed with black. DT ringed with black. DT on segments 2-10
orange to yellow. DT on segment 1 blue. CT yellow with ring of black dots. Other
tubercles blue.

Fifth instar: Green, lighter dorsally. Green lobes, yellow frons, blue to grey
clypeus, grey labrum, blue palpi. Spiracles white ringed with black. DT on seg-
ments 2-4 orange with black ring. DT and ULT on segment 1 blue to black rudi-
mentary dots. Other tubercles white with a black ring, slightly bluish above ring.

Summary

The quadruple hybrid [(H. cecropia ¢ x H. gloveri 2?) & xX H. rubra
*|é x If. columbia 2 was reared successfully. Willow and wild cherry
were excellent foodplants and survival was excellent at all stages. Rep-
resentative specimens have been donated to the American Museum of

|

History.

ral

VOLUME 25, NUMBER 1 1c

Acknowledgments

I would like to express my thanks to Mr. Cyril F. dos Passos for re-
viewing the manuscript and making valuable suggestions, and to Mr.
Frederick W. Case II and Mr. John T. Sorensen for donating H. columbia
pupae, without which this project could not have been completed.

Literature Cited

Coxtiins, M. M. AnD R. D. Weasr. 1961. Wild Silk Moths of the United States,
Collins Radio Corp.

Crorcu, W. J. B. 1956. A Silkmoth Rearer’s Handbook. The Amateur Entomolo-
gists Society.

Prippite, T. R. 1966. Techniques for reducing mortality when rearing larvae of
the cecropia moth (Saturniidae). Jour. Lepid. Soc. 20: 119-121.

SWEADNER, W. R. 1937. Hybridization and phylogeny of the genus Platysamia.
Annals of the Carnegie Museum 25: 163-242.

WiuiaMs, C. M. 1955. Physiology of insect diapause. X. An endocrine mecha-
nism for the influence of temperature on the diapausing pupa of the cecropia
silkworm. Biol. Bull. 110: 210-218.

SOME NOTES ON THE PAPILIONIDAE OF
MANUS ISLAND, NEW GUINEA

James A. EBNER

Okauchee, Wisconsin

Manus Island is a relatively obscure island which anchors the western-
most chain of the Bismark Archipelago. It is approximately 150 miles in
length and the principal island of the Admiralty Island group. The near-
est New Guinea mainland lies some 300 miles south, and Papuan in-
fluence is reflected in the fauna of Papilionidae of Manus.

In the past, few specimens have come into the hands of collectors from
this remote island, largely due to the hostile natives that once inhabited
the land. Also Manus was not situated along major shipping trade lanes,
and consequently few opportunities existed for visits by travelers. Web-
ster in 1897 was apparently the first to collect the nearly inaccessible is-
land, but was forced to depart hastily because of the unfriendly natives.
He succeeded in gathering some papilionid species, however, and these
were included in Seitz’s treatment of Indo-Australian butterflies.

The A. S. Meek expedition to the Admiralty Islands and adjacent lo-
calities in 1913 was more successful than Captain Webster's. Meek,
commissioned by Lord Rothschild of the Tring Museum in England, was
able to collect Manus Island during September and October, and to assess

74 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

the island’s fauna accurately. When Lord Rothschild received Meek’s
material a report of the species taken appeared in “Novitates Zoologicae,”
XXII, in 1915. This publication recorded eight papilionid species for
Manus of which four were described as new subspecies and another,
Papilio cartereti Oberthur (= Papilio weymeri Niepelt in Strand) was
an apparent new discovery deserving species status.

Apart from the collections made by Webster, Meek and a few others,
the Admiralty Island group continued to be an essentially by-passed loca-
tion for many years. Fortunately the author was successful in obtaining
the services of a resident collector on Manus in 1968. Papilionidae were
supplied at regular intervals for more than a year, and this more recent
material substantiates the previous works of Webster and Meek. The an-
notated list of Papilionidae is herein offered to consolidate and supple-
ment information concerning the Papilionidae of the Admiralty Islands.
The arrangement of species follows that of Munroe (1961).

Graphium codrus auratus (Rothschild )
Papilio codrus auratus Rothschild, 1898, p. 218.

Described from 229, St. Gabriel, Admiralty Island, February, 1897 (Captain
H. C. Webster). Rothschild (1915, p. 194) 2¢ 6 and 299 from Manus (Meek
Expedition ).

The rarest known Papilio on Manus. Difficult to net and disdainful of
lures. December to March generally favorable flight months. No varia-
tion in the few examples received. Distinguished from other codrus sub-
species by the presence of nine bright yellow spots on each forewing.

Graphium macfarlanei admiralia (Rothschild)
Papilio macfarlanei admiralia Rothschild, 1915, p. 195.
Described from 72 9, Manus Island (Meek Expedition ).

specially common to abundant in April and May, but occurring all
year round. Sex ratio of adults seemingly equal. This subspecies char-

=>

Figs. 1-11. Manus Island Papilionidae (full expanse in brackets). 1, Graphium
agamemnon admiralis Rothschild @ , Oct. 1968, (68 mm); 2, G. macfarlanei admiralis

Rothschild §, Jan. 1969, (74 mm); 3, G. macfarlanei admiralia Rothschild OF an
1969. (70 mm); 4, Papilio cartereti Oberthur é, May, 1969, (1S Sinn eee.
cartereti Oberthur 9, May, 1969, (125 mm); 6, P. ulysses gabrielis Rothschild 2,

June, 1968, (102 mm); 7, P. ulysses gabrielis Rothschild 2, Oct. 1968, (110 mm);
8, G. codrus auratus Rothschild 9, Dec., 1968, (77 mm); 9, P. polydorus manus
Talbot 3, April, 1968, (83 mm); 10, P. polydorus manus Talbot &@ melanic, Dec.,
1968, (72 mm); 11, P. polydorus manus Rothschild ©; Dec:, 1968 (85 imme

VOLUME 25, NuMBER 1

76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bi

ie

Figs. 12-15. Manus Island Papilionidae. 12, Papilio phestus reductus Rothschild
6, Oct., 1968, (76 mm); 13, P. phestus reductus Rothschild 9, July, 1968, (68
mm); 14, Ornithoptera priamus admiralitatis Rothschild ¢, April, 1969, (140 mm);
15, O. priamus admiralitatis Rothschild ¢@, April, 1969, (157 mm).

acterized principally by the prominent black margins of both fore- and
hind wings. Primaries with greatly reduced green submarginal spots,
sometimes barely indicated against the black ground color. Hind wing
of the male lacking submarginal band entirely or displaying but a trace
of this band in the form of two or three faint spots. Hind wing of the

female with a greater area of black ground color, resembling seminigra
sutler from New Britain.

Graphium agamemnon admiralis (Rothschild )
Papilio agamemnon admiralis Rothschild, 1915, p. 195.
Described from 3¢ 6, 59 9, Manus Island (Meek Expedition).

Not common in general. Male to female ratio approximately 4 to 3.
No appreciable variation. The subspecies characterized in both sexes by

the prominent and broad black margins on the upper surfaces. Submar-

VoLuME 25, NuMBER 1 on

ginal spots very greatly reduced, sometimes almost absent. Hind wing
resembling that of P. agamemnon argynnus ( Druce).

Papilio cartereti Oberthur

Papilio cartereti Oberthur, 1914, p. 187 (April 8).

Papilio weymeri Niepelt in Strand, 1914, p. 53; plate 11, fig. 1 (Jan? May?).

Papilio cartereti Oberthur (=P. weymeri); Oberthur 1915, p. 499; plate 311, fig.
4602; plate 312, fig. 4603.

Papilio weymeri Niepelt; Rothschild, 1915, p. 194.

Papilio weymeri Niepelt in Strand; Strand 1926, p. 410.

Papilio weymeri Niepelt in Strand; Bryk 1930a, p. 192.

Papilio weymeri Niepelt in Strand (= P. cartereti Oberthur); Bryk 1930b, p. 608.

Papilio ? weymeri Niepelt; Munroe 1961, p. 43.

P. cartereti described from 1é¢ and 192 from the Admiralty Islands collected by
the Meek Expedition; named by Oberthur after the navigator Carteret who appar-
ently visited the islands in 1767. P. weymeri described from a single female from
the Admiralty Islands (not collected by Meek; see Rothschild 1915, p. 192). Roths-
child (1915, p. 194) recorded 66 6 and 62 2 from Manus (Meek Expedition).

It has not so far proved possible to establish beyond doubt the priority
of the synonymous names Papilio cartereti Oberthur and Papilio weymeri
Niepelt in Strand, both names without doubt having appeared in the first
six months of 1914. Reference to page 528 of Bull. Soc. ent. Fr. for 1914
establishes the date for P. cartereti as 8th April 1914 (the date of publica-
tion for “Seance 25 Mars” in which Oberthur’s description appeared).
Part 1 of Lepidoptera Niepeltiana in which P. weymeri was described by
Niepelt has the forward dated October, 1913, but the paper cover is
clearly dated 1914. Copies in the libraries of the British Museum ( Nat-
ural History) and the Royal Entomological Society of London give little
further information. The copy formerly in the Zoological Museum, Tring
(The Museum of Walter Rothschild), however, has been overprinted
“Januar” and has a Tring Museum date stamp “Received 25 July 1914.”
Oberthur (1915, p. 499) states that his own description of the species
appeared in March and thus a little earlier (“un peu avant lui’) than
Niepelt’s, which he claims appeared in May (“paru en Mai’). It would
seem that there was some delay in the distribution of Niepelt’s privately
published work; the British Museum (Natural History) did not receive
a copy until 22nd May 1920 and the Royal Entomological Society did
not obtain a copy until it purchased part of the Joicey Library (the copy
in question unfortunately having no original receiving date on it). How-
ever, it is known that the description of P. cartereti appeared on 8th April
1914 while that of P. weymeri appeared certainly sometime before 25th
July 1914. Whether one accepts the single overprinting date of January
on the ex Tring copy of Lepidoptera Niepeltiana or Oberthur’s claim

78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

that it appeared in May is at present a matter of opinion. The present
location of the holotype of P. weymeri is not known to the author; two
syntypes of P. cartereti are in the British Museum (Natural History).

Only member of the Papilionidae confined soley to Manus Island.
Males flying erratically and never seeming to settle. Females less com-
mon than males; flying slowly in and among small, stunted brush. June,
July and August exceptionally good flight months.

Male greatly resembling P. bridgei Math. In cartereti ground color
very dark brownish-black. Upper surface of the forewing always with a
band of eight cream colored spots. Band of hind wing broad and uni-
formly greyer than spots of forewing. Variation in males almost non-
existent except in size. Female with a generally browner ground color
on upper surface than male. Medium band of forewing dusted, smoky
light brown. On hind wing a large postmedian band of six brilliant blue
patches, terminating at the inner margin with an additional bright orange
spot. Submarginal crescents pale orange.

Papilio phestus reductus Rothschild
Papilio phestus reductus Rothschild, 1915, p. 193.

Described from a “type” (holotype) and a further pair (paratypes) from Manus,
collected by the Meek Exvedition.

Not especially common on Manus. Males slightly more prevalent than
females. According to Rothschild in his original description of reductus,
this form is distinguishable in male by lack of any light spot in cell of
hind wing. This characteristic, however, also prevalent in males from
New Ireland. Prominent deep-orange patch at the inner margin of Manus
Island phestus always triangular or nearly so. New Ireland phestus with
same spot squared or rectangular. In female of reductus a greater varia-
tion in the number of white and cinnamon spots on hind wing.

Papilio ulysses gabrielis Rothschild
Papilio ulysses gabriclis Rothschild, 1898; p: 207,

Described from a series of 64 and one 2 from St. Gabriel, Admiralty Islands,
collected by Captain H. C. Webster, February, 1897. Rothschild (1915, p. 194 re-
ceived a further 92 ¢ from Manus (Meek Expedition ).

Prevalent the year round and quite common. As many as 15 adults
can be netted in an hour, the majority imperfect in condition. Females
considerably rarer. Extent of blue in both sexes varying noticeably, but

always remaining greatly reduced when contrasted with other ulysses
subspecies.

VoLUME 25, NuMBER 1 79

Pachlioptera polydorus manus (Talbot)
Papilio polydorus manus Talbot, 1932, p. 155.
Papilio polydorus utuanensis Ribbe; Rothschild 1915, p. 193.

Described from 2¢ ¢ and 59 @ from Manus, Sept.-Oct., 1913 (presumably col-
lected by the Meek expedition). Rothschild (1915, p. 193) recorded 76 6 and
62 2 from Manus collected by Meek, under the name P. polydorus utuanensis.

Common, tending to be abundant periodically. Male-female ratio
nearly identical. Some males melanistic, displaying dark suffusion over
more normal, creamy white spots of hind wing, thereby approaching P.
polydorus dampieriensis Hag.

Ornithoptera priamus admiralitatis (Rothschild )

Papilio priamus admiralitatis Rothschild, 1915, p. 192.
Described from 1¢ and 82 2, Manus (Meek Expedition).

Common and widespread, flying throughout the year. Approximately
an equal ratio of sexes in the wild. In male, admiralitatis totally lacking
dorsal green scaling on veins in disc of forewing. On hind wing, above,
green scaling denser than in O. priamus bornemanni Pagenst, its nearest
ally. O. p. admiralitatis males rarely lacking golden yellow patch in
costal area. In female, according to Rothschild, admiralitatis distinguish-
able from bornemanni by shorter, rounder wings and _blackish-grey,
brighter ground color. Furthermore, pale patches of hind wing extend-
ing almost to cell; some specimens with a white spot in cell. On hind
wing dark spots in light patches very much larger than in bornemanni.

Acknowledgments

The author is sincerely indebted to the painstaking work of Mr. R. I.
Vane-Wright of the British Museum (Natural History). His efforts con-
cerning the research of the literature has enabled the author to document
this article with comparatively rare and pertinent material. In addition,
Mr. Vane-Wright has assisted greatly in the development of the final
manuscript and certainly without his help this article could not have
been completed.

Literature Cited

Bryk, F. 1930a. Papilionidae II (Papilio). Lepid. Cat. 37.

1930b. Papilionidae III. Lepid Cat. 39.

Munroe E. 1961. The Classification of the Papilionidae (Lepidoptera). Can.
Ent. Suppl. 17.

Nizpett, W. 1914. In Strand, E. Lepidoptera Niepeltiana. 1.

OseRTHUR, C. 1914. Description d’une nouvelle espéce de Papilio des iles de
YAmirauté (Lep. Rhopalocera). Bull. Soc. Ent. Fr. (1914) pp. 187-188.

-—— 1915. Etudes de Lépidoptérologie Comparée 10. Rennes.

80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

RorHscHitp, W. 1898. Some New Lepidoptera from the East. Novit. zool. 5:
216-219.

1915. On the Lepidoptera in the Tring Museum sent by Mr. A. S. Meek
from the Admiralty Islands, Dampier, and Vulcan Islands. Novit. zool. 22:
192-208, 387-402, etc.

Serrz, A. 1927. Macrolep. World. 9, The Indo-Australian Rhopalocera. Stutt-

art.

sen E. 1926. Liste des Rhopalocéres et Grypocéres exotiques décrits dans mes
travaux jusquen 1926. Bull. Soc. Zool. Fr. 51: 397-418.

Talbot, G. 1932. New Forms of Lepidoptera from the Oriental Region. Bull. Hill
Mus. Witley 4: 155-169.

A NOTE ON “MATING FLIGHT OF BUTTERFLIES WITH
MIMETIC FEMALES AND NON-MIMETIC MALES”

My friend R. H. Carcasson’s note under the above heading (1970, Journ. Lep.
Soc., 24: 72) caused me to look up my own records; the following may be of in-
terest:

Belenois thysa Hpffr. (Pieridae )—Nyali, 4.vii.70. Male flew.

Acraea encedon 1. (Acraeidae )—Kampala, 8.v.49 and 30.vii.49 (two pairs). Fe-
male flew.

Euryphene mardania orientis Karsch (Nymphalidae )—Shimba Hills, 31.v.70. Fe-
male flew.

Precis clelia Cr. (Nymphalidae )—Kampala, 8.v.49. Female flew.

Hypolimnas misippus L. (Nymphalidae )—Kampala, 8.v.49. Female flew.

Castalius calice Hpffr. (Lycaenidae )—Nyali, 9.vii.70. Female flew.

Both sexes of A. encedon, the female of H. misippus and the female of E. mar-
dania mimic Danaus chrysippus L. P. clelia and C. calice are not mimetic and both
sexes of B. thysa might be said to mimic Mylothris.

My own feeling is that the flying position of mated pairs is more of a family, or
possibly subfamily, characteristic, and has no direct relationship with mimicry.

D. G. SEvastoputo, P. O. Box 5026, Mombasa, Kenya.

SOME RECORDS OF EURISTRYMON ONTARIO (LYCAENIDAE)

Euristrymon ontario ontario Edwards is rare enough in eastern North America
that any captures deserve to be put on record, particularly when the associated en-
vironmental circumstances also can be given.

On the basis of what little habitat information I had, the shale barrens of the mid-
Appalachians seemed to be an appropriate place for this little-known species. An
opportunity to look for it there came in June 1968 when my wife and I drove from
Mlorida to Pittsburgh. We planned our route to cross the Appalachians in Virginia
at a point where shale barrens were known to occur, and on 14 June we stopped in
the late afternoon to collect in a typical barrens area in Alleghany County, Virginia,

ar ¢ lifton Forge. Between 4 and 6 PM EDST I took five nearly fresh specimens

ttaTrwo.

vere on the newly opening flowers of Dogbane (Apocynum cannabinum)

VoLUME 25, NuMBER 1 81

which grew in a large, dense stand on a steep, east-facing slope of road fill, the rays
of the late afternoon sun just grazing the plants. Only a few feet away was a low,
open forest of hard pine and oaks. One of the latter, Quercus ilicifolia (or marilan-
dica), is a low shrubby species that may possibly be the larval foodplant of ontario.
Also taken at the same time and place were: Satyrium calanus falacer Godart (22,
fresh), S. liparops strigosum Harris (16, fresh), both taken on Dogbane; Achalarus
lyciades Geyer (fresh) and Thorbyes (apparently both pylades Scudder and bathyllus
Smith), on the flowers of Vipers Bugloss (Echium vulgare); Epargyreus clarus
Cramer and Speyeria cybele Fabricius, both on a single plant of Common Milkweed
(Asclepias syriaca) growing among the Dogbane. A pair of rather worn Hesperia
sassacus Harris was also taken.

My favorite shale barrens area is in Green Ridge State Forest, Allegany County,
Maryland. Rumor has long had it that ontario occurs here, and the general habitat is
similar to the Virigina locality. So as soon as possible after we reached Pittsburgh,
and armed with my newly augmented knowledge of the environmental choice of
ontario, my wife and I headed for Green Ridge and spent a day (22 June 1968)
looking for it, but absolutely in vain.

The next year, 1969, I was unable to visit Green Ridge at the right time, but in
1970 I went there again and spent 19-20 June combing the area carefully over a wide
range of possible habitats. On the 19th I found none at all, even though several of
the places searched seemed ideal. But on the 20th my luck changed: in three dif-
ferent locations I took a total of four specimens, all rather worn.

These captures were as follows. (a) “Boy Scout Meadow” [my term]: one each
in two different stands of Dogbane, a moderate-sized stand at least 50 feet from the
forest, and a large stand immediately adjoining the forest; the forest is low and
open, dominantly hard pine with oak and hawthorn admixed, in the valley of
Fifteen-Mile Creek (elevation 790 ft.); (b) Sugar Bottom Road: one on the white
flowers of Wild Quinine (Parthenium integrifolium) along the roadside at the edge
of a tall, fairly dense forest of mixed pine, oak and probably hickory along the crest
of a ridge (elevation 940 ft.); on the other side of the road was a large cut-over
area of stumps, low shrubs and forbs; (c) White Sulphur Community Pond: one
on flowers of Dogbane growing in a moderate-sized stand about 25 feet from the
edge of a low forest of pine and oak (elevation 750 ft.). Quercus ilicifolia is a com-
mon species in the Green Ridge area and formed part of the forest in several of the
above areas, perhaps all of them. In addition to ontario, 28 species of butterflies
were taken during these two days. The more significant of these are: Limenitis
arthemis astyanax Fabricius (the commonest butterfly at the time); Epargyreus
clarus (common, on flowers of both Dogbane and Viper’s Bugloss); Speyeria cybele
(rather common, on flowers of Dogbane and Butterfly Weed, Asclepias tuberosa);
Achalarus lyciades (one only); Satyriwm calanus falacer and S. liparops strigosum
(both somewhat worn, on Dogbane, falacer much the commoner and also on leaves
at woods edges); Chlosyne nycteis Doubleday (fairly common and fresh); Thorybes
pylades (worn). Lethe portlandia anthedon A. H. Clark was just beginning to ap-
pear (only a single specimen was seen), as was Pompeius verna Edwards.

Our preparator, Mr. John Bauer, has long been interested in ontario, and after I
had brought back the Maryland specimens he went out to try his luck, despite the
already late date. On 28 June, the first day with favorable weather, he went west of
Pittsburgh and at a spot near the common boundary point of Allegheny, Beaver and
Washington Counties he took a single, much worn specimen, a new species record
for Pennsylvania. Western Pennsylvania is an area of essentially deciduous forests
of various kinds, and this locality is in a small area of mixed maple and White Oak
forest; it is quite different from the Virginia and Maryland localities. His single
specimen was taken on the flowers of Common Milkweed. At the same time and
place Mr. Bauer also took Satyriuwm calanus falacer (common, somewhat worn, on

82 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

leaves), S. liparops strigosum (somewhat worn) and Harkenclenus titus Fabricius
(fresh) (the latter two on Butterfly Weed), a single female of S. acadica Edwards
(on milkweed), along with Chlosyne nycteis (fresh), Polites peckius Kirby and
Pompeius verna (both fresh), Lethe portlandia (fresh) and Thorybes pylades (very
worn ).

Sa several records of ontario permit some generalizations on habitat and flight
period that may be of help to others in searching for it. Proximity to a forest, pref-
erably low and rather open, of hard pine (Pinus virginiana is probably the species
most often involved) and oak (Quercus ilicifolia presumably one of them) is a
common factor for most of the records. So, too, is a nearby open meadow, or other
suitable open area, where Dogbane grows. Dogbane seems to be by far the most
favored food flower, although single captures in Green Ridge and in Pennsylvania
indicate that other flowers may occasionally be visited. Dr. C. F. dos Passos informs
me that Dogbane is the favored food flower of ontario in New Jersey, and Professor
Ernest M. Shull makes the same observation for northern Indiana.

In view of the condition of the specimens and their dates of capture I would
estimate that normally the flight period of ontario in the Virginia-Pennsylvania re-
gion begins on the 10th to 15th of June (possibly a little earlier) and lasts no more
than about 20 days, an exceptionally short flight period for a butterfly. The 15th-
20th of June seems to be about the optimum time for seeking it in reasonable num-
bers and in good condition. It appears just about when Hesperia sassacus is ending
its flight, perhaps about 10 days after Thorybes pylades has begun to fly, and just
before the appearance of Speyeria cybele, Satyrium falacer, S. liparops and Lethe
portlandia.

E. ontario is always uncommon, and usually rare. Perched on Dogbane flowers
it is usually not reliably distinguishable from falacer or liparops which occur with it.
In the Green Ridge area I searched approximately 15 moderate to large-sized stands
of Dogbane and found ontario in only three. In its daily behavior it may resemble
falacer in making use of the forest for roosting at night, and perhaps for mating, but
feeding much of the day at flowers usually in nearby fields or other open areas.

The records discussed above may be summarized as follows:

Virginia: Alleghany Co.: 4 mi E of Clifton Forge, on Va. 42 about % mi N of ject.
with US 60; 14.vi.1968, 3 ¢@ 29 (H. Clench).

Maryland: Allegany Co.: Green Ridge State Forest, 6 mi E of Flintstone, 20.vi.
1970 (H. Clench): (a) “Boy Scout Meadow” on Fifteen-Mile Creek Rd. about
1.5 mi N of US 40, 1¢ 19; (b) Sugar Bottom Road, ca. 1 mi S of US 40, 19; (c)
White Sulphur Community Pond, ca. 2 mi S of US 40, 19.

Pennsylvania: Allegheny Co.: Murdocksville, ca. 3 mi WNW of Clinton, about at

common point of Allegheny, Beaver and Washington Cos., 28.vi.1970, 13 (John
Bauer ).

Harry K. CLENcH, Carnegie Museum, Pittsburgh, Penna.

SHAPIRO COLLECTION AT CORNELL

The Arthur M. Shapiro collection of about 13,000 mounted and determined

Rhopalocera has been placed in the Department of Entomology and Limnology at
Cornell University. It includes 9436 specimens (108 species) from upstate New
York and Pennsylvania, of which highlights include long series of all the northeastern

Lethe (including the newly recognized species, L. appalachia), Limenitis arthemis/
astyanax intergrades, representatives of the newly discovered Lycaeides melissa sam-
uclis population from western New York, the unique McLean Bog, N. Y. population of

VoLUME 25, NuMBER 1 83

Poanes massasoit, and such little-known skippers as Euphyes dion and bimacula,
Poanes viator, and Hesperia leonardus and metea. Shorter series of special interest
are Coenonympha tullia heinemani, a unique eastern Adirondack Cercyonis, Erora
laeta, Incisalia spp., and Colias interior from Pennsylvania and Tug Hill, N. Y.
Singletons include a hybrid of Limenitis arthemis and archippus and a gynandromorph
of Pieris rapae, both taken wild in Tompkins Co., N. Y.

The British series includes 1393 specimens (24 species), of which 558 are of the
very strange Pieris napi-group taxon from northern Scotland (about 20 localities).
There are far-northern Scottish Lycaena phlaeas, Polyommatus icarus, and Coeno-
nympha pamphilus, and a bred gynandromorph of English P. napi. Thymelicus
lineola is represented by examples from both countries, including 116 documenting
the spread of this introduced species through central New York since 1968.

Among 1026 butterflies in miscellaneous series are over 200 of a Colias philodice-
eurytheme hybrid swarm in southeastern Arizona, various mutants of the eastern
Colias, and a series of Lycaenopsis pseudargiolus from the New Jersey pine barrens
including extreme lucia. Finally, there are 1111 bred Pieridae illustrating genetic
and environmental experiments on color and pattern regulation.

L. L. PecuuMan, Cornell University, Ithaca, N.Y.

SPECIMENS DAMAGED BY CARPENTER ANTS

Even the unseasoned collector is aware of the necessity for protecting specimens
in storage or cabinet with paradichlorobenzene or naphthalene, to guard against
dermestid infestation. In any closed container, these substances repel all injurious
pests. When used in a drying cabinet, specimens on setting boards can also be pro-
tected.

When setting boards are not enclosed, however, specimens may be subject to one
uncommon pest, the Carpenter Ant, Camponotus pennsylvanicus. On one occasion I
found my boards swarming with workers of pennsylvanicus, and the abdomens of
several specimens had been eaten. The setting board grooves were stained yellow
beneath the specimens with what may have been a formic acid solution, secreted by
the ants to soften the abdomen tissues while feeding.

Since dermestid damage is a slow process, there is little danger of destruction dur-
ing the relatively short time specimens are on the setting boards. Damage from C.
pennsylvanicus, however, can occur in only minutes. To prevent its recurrence, I
treated the edges of all setting boards with a commercial insecticide having strong
residual properties. No further ant damage has been noted since this precaution was
taken.

J. B. Woop, 140 Pines Drive, Henderson, Kentucky.

IRWIN COLLECTION TO ILLINOIS NATURAL HISTORY SURVEY

The writer of this note has donated the bulk of his collection to the Illinois Nat-
ural History Survey, Urbana, Illinois. It consists of approximately 3,800 pinned
specimens of mostly North American butterflies, with emphasis on the Illinois fauna.
A number of species contained in the collection were not previously represented in

84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the Survey collection by Illinois specimens; four species are represented by the only
Illinois specimens known.

Also included are 63 specimens originally in the collection of William Henry Ed-
wards and bearing his holograph labels; of these one is an Edwards syntype. This
material was listed and its history discussed in a paper in this Journal (Irwin, 1966,
J. Lepid. Soc. 20: 156-162). It has been extensively studied by F. Martin Brown
during his current researches on the Edwards butterfly types (see Brown, 1964,
Trans. Amer. Ent. Soc. 90: 323-413, and subsequent papers in this series).

With the addition of the Irwin collection, that of the Survey becomes the largest
and most complete of Illinois butterflies in existence. The Survey is second only to
the Field Museum among Illinois institutions in total holdings of Lepidoptera. The
writer is currently incorporating several other individual collections as well as his
own into the unified Survey butterfly collection, while the remainder of the Lepi-
doptera is being curated by Dr. Robert W. Poole of the Survey staff.

The writer plans to continue to collect and study Lepidoptera, and to complete
the faunal survey of Illinois butterflies which he and Dr. John C. Downey have been
conducting for several years.

Roperick R. Irwin, 24 East 99th Place, Chicago, Illinois.

BOLORIA EUNOMIA LADDI (NYMPHALIDAE) IN COLORADO

On 5 July 1967, John Sorensen of Waterloo, Iowa, Pat Conway of Chicago and I
collected a small series of Boloria eunomia (Esper) in a willow bog just above
10,000+ feet in Gunnison County, Colorado. After examination, these specimens
appear to belong to the “Wyoming” subspecies, laddi (Klots), rather than to the
“Colorado” subspecies, caelestis (Hemming). They agree with the laddi phenotype
by having dark, red-brown coloration on the basal area of the underside of the hind
wing and on the subapical patch on the underside of the forewing; rather than the
light, yellow-brown coloration of caelestis. Their appearance on the upperside does
not differ from either laddi or caelestis. I have no hesitation in assigning them to
laddi, even though they are separated from Wyoming populations of this subspecies
by considerable distance.

Other than a record by Scott Ellis (19 July 1964, also Gunnison County), these
are the only known examples of eunomia from west of the continental divide in Colo-
rado. I suspect that the laddi phenotype will be found to occur along the western

slope in Colorado, while the caelestis phenotype will be endemic to the Colorado
eastern slope.

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin.

FLIGHT PATTERN OF THE MALE OF ANISOTA VIRGINIENSIS
(CITHERONIIDAE )
a. colony of fourth-instar larvae of Anisota virginiensis (Drury) was found feed-
ing on Water Oak, Quercus nigra (L.), early in July, 1970, about seven miles north
of McClellanville, South Carolina

The larvae were reared successfully to pupation.
cc were preserved by both inflation and vacuum
Che authors were especially interested in rearing this species through

About half a dozen mature larvae
freeze-drying.

VoLUME 25, NuMBER 1 85

to the adult stage for, contrary to the usual pattern, their collection contains a
series of females all caught in light traps, but there is not a male in the crowd. A
search of the literature working backward from Lutz and Holland through Seitz,
Packard, McDunnough, Forbes, etc., revealed a multitude of descriptions and lots
of lovely plates and figures of the males and females, but nowhere an answer to
the evident dislike of males for our light traps. It was a chance question during a visit
by Drs. D. C. Ferguson and J. G. Franclemont that began to shed some light on the
mystery.

“Aha,” they said in unison. “It’s no wonder. Males of virginiensis are crepuscular
and don’t fly after dark!” And so we waited while our twenty-one pupae ticked
along. On 21 July a female emerged early in the morning and within one week
the whole lot had proved viable and had emerged; there were about equal num-
bers of males and females. Without exception both sexes emerged soon after sun-
rise and were expanded and ready for flight by about 9 AM, EST. They were in
a screen wire cage on a sheltered porch outdoors, and we killed and mounted all the
males as soon as they were dry enough (except for one which escaped with re-
markable rapidity). The females, which remained quite docile, were left alive in
the cage in hopes that they would attract wild males in the late afternoon and thus
prove the crepuscular theory. In vain we sat and watched until it was too dark to
see, and again the following mid-afternoon till dark. The next morning about eleven
o clock one of us (CRE) happened to pass by the cage and in utter astonishment
saw a very swift red object buzzing rapidly about. A hasty grab for a handy net,
and a wild male virginiensis was bagged. The following day about noon one of our
wives who maintains a close interest in our odd activities reported the same phe-
nomenon. She likewise grabbed the nearby net but failed in the capture. There-
after we ceased our fruitless afternoon and evening vigils and instead stood watch
from sunrise on. In three days, before our last female died her natural death, we
caught thirteen more wild males, all between 10 AM and 1:30 PM, EST. They
fly rapidly and erratically, reminding one of fast skippers or small Sphingidae,
often stopping to hover nearby for a moment and then darting off again. Patience
is required of the stalker; wild mid-air sweeps of the net are generally disasterous,
but the temptation tantalizing. Thus the actual number of males attracted may
have exceeded those captured. The wild males we did capture were almost without
exception fresh and in good condition. We did not mate any of the females, and
found their natural infertile adult life to be about five days.

One strange and so far unexplained incident occurred. On two occasions a fly-
ing object paid a swift passing visit to the caged females. We should have be-
lieved it to be virginiensis by size, speed and flight pattern but it was dark in color,
not the typical deep red of virginiensis which is very conspicuous in flight. The
first of these UFO’s zipped away too fast for identification, but it bore resemblance
to the second which hovered at the cage just long enough for positive identification
as Amphion nessus (Cram). While the latter is a common day-flier in this area, it
certainly bears no close relationship to the Citheronidae, and its attraction seems a
mystery.

RicHArpD B. DominicK AND CHARLES R. Epwarps, The Charleston Museum,
Charleston, South Carolina.

86 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

BOOK REVIEW

MARIPOSAS DE VENEZUELA. By Michael Schmid and Bradford M. Endicott. 1968.
xi + 67pp., 142 color illus. Publisher: L. Levison Junr., Copenhagen. [available in
U.S. from Entomological Reprint Specialists, P.O.Box 207, East Lansing, Michigan;
price $9.95]

This, the first book ever to treat Venezuelan butterflies, is, as it should be, in
Spanish; but, fortunately for the North American reader, contains a complete Eng-
lish text in the back. The 142 colored plates are excellent. Those species depicted
were undoubtedly chosen for their beauty, but still comprise a characteristic repre-
sentation of the Venezuelan fauna. All but a very few, e.g. Morpho species and
Agrias narcissus, are species that occur in the populated Venezuelan coastal plain.

The commentary on each species is brief but interesting and easily read. The
English text is not an exact translation of the Spanish but in a few places has been
adjusted slightly for an English audience as, for instance, in the addition of a few
English vernacular names. It is easy to fault the book for the outdated nomenclature
used, most of which dates back to Seitz. I found 38 of 113 butterflies placed in a
taxonomically incorrect genus. A table of “recent name changes” in the back of the
book corrects ten of these, but another seven are changed to still another improper
usage.

This book is an introduction to Venezuelan butterflies and not a definitive study
of them. For the Venezuelan with a casual interest in Lepidoptera or for the North
American Lepidopterist with a casual interest in Venezuela, it has a definite place.

Joun H. Masters, P.O. Box 7511, St. Paul, Minnesota.

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
for material in excess of 20 printed pages, at the rate of $17.50 per page. Address
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Contributors should prepare manuscripts according to the following instructions;
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Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
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full scientific name, with authors of zoological names. Underline only where italics
are intended in the text (never in headings). References to footnotes should be
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Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
heading LireratureE Cirep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: Al! photographs and drawings should be mounted on stiff, white
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Tables: Tables should be numbered consecutively in Arabic numerals. Headings
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Material not intended for permanent record, such as current events and notices,
should be sent to the editor of the News: Dr. C. V. Covell, Dept. of Biology, Univer-
sity of Louisville, Louisville, Kentucky 40208.

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyrit F. pos Passos
Price, postpaid: Society members—$4.50, others—$6.00; uncut,
unbound signatures available for interleaving and private binding,

same prices; hard cover bound, add $1.50. Revised lists of the
Melitaeinae and Lycaenidae will be distributed to purchasers free.

ALLEN PRESS, INC. eRinTeD LAWRENCE, KANSAS
usr

CONTENTS

Bowden, S. R. American white butterflies (Pieridae) and English food-

Plants cc 6-12
Brussard, P. F. Field techniques for investigations of population structure

in a “ubiquitous” butterfly 2000) 240) 22-29
Clench, H. K. Some records of Euristrymon ontario (Lycaenidae) —_.____ 80-82
Dominick, R. B. and C. R. Edwards. Flight pattern of male of Anisota

virginiensis (Citheroniidae) 0... 84-85

Ebner, J. A. Some notes on the Papilionidae of Manus Island, New Guinea 73-80

Emmel, T. C. Symbiotic relationship of an Ecuadorian skipper (Hesperi-
idae) and Mazxillaria orchids

Ferris, C. D. and M. Fisher. A revision of Speyeria nokomis (Nymphal-

Robe) te GN a 4-52
Gray, R. E. Papering Lepidoptera in glassine envelopes __.......---- 65-68
Hardwick, D. F. The life history of Heliothis oregonica (Noctuidae) 1- 6
Irwin, R. R. Irwin collection to Illinois Natural History Survey 83-84
Kendall, Roy O. and C. A. Kendall. Lepidoptera in the unpublished field

notes of Howard George Lacey, naturalist (1856-1929) _.....- 29-44
Lindsay, S. A. A host plant for northern populations of Euchloe olympia

(Pieridae ) i030) 0 ali) Nr 64
Masters, J. A. Book Review: Mariposas de Venezuela __........ 86
Masters, J. A. Consul panariste (Nymphalidae) in Venezuela __....__ 19
Masters, J. A. Boloria eunomia laddi (Nymphalidae) in Colorado _.__.___ 84
Miller, L. D. and T. C. Emmel. The Brazilian “Cercyonis” (Satyridae) 12-19
Pechuman, L. L. Shapiro Collection at Cornell 82-83
Sevastopulo, D. G. A note on “Mating flight of butterflies with mimetic

females and non-mimetic males”) 00 ee 80
Stary, P. and I. K. Kaddou. Observations on the biology of Ocnerogyia

amanda Stgr. (Lymantriidae), a pest of Ficus in Iraq 53-57
Straatman, R. The life history of Ornithoptera alexandrae Rothschild ___ 58-64
Wood, J. B. Specimens damaged by carpenter ants 83

Wright, D. A. Hybrids among species of Hyalophora 68-73

BY | Volume 25 1971 Number 2

JOURNAL

) of the

_ LEPIDOPTERISTS’ SOCIETY

one.
~

BY, Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

i
a,
MY
+

4
Ay
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4
; 19 May 1971
;
;

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

D. F. Harpwicx, Editor of the Journal
C. V. CovEtx, Editor of the News
S. A. Hessen, Manager of the Memoirs

EXECUTIVE COUNCIL

C. L. Remincton (New Haven, Conn.) President

Lioyp M. Martin (Prescott, Ariz.) President-elect

H. A. FrEEMAN (Garland, Texas) 1st Vice President
JuL1an Jumaton (Cebu City, Philippines) Vice President
K. W. Pure (Fairbanks, Alaska) Vice President

S. S. Niconay (Virginia Beach, Va.) Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

LzE D. Miter (Sarasota, Fla.) Secretary-elect

Members at large (three year term): E. C. Wetuine (Merida, Mexico) 1972
A. E. Brower (Augusta, Me.) 1971 ANDRE BLANCHARD (Houston, Texas) 1973
W. C. McGurrin (Ottawa, Ont.) 1971 R. B. Dominick (McClellanville, S. C.)
Y. NExRuTENKO (Kiev, U.S.S.R.) 1971 1973
B. Matuer (Clinton, Miss.) 1972 J. P. Donanut (Los Angeles, Calif.)
M. Ocata (Osaka, Japan) 1972 1973

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
and the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

Membership in the Society is open to all persons interested in the study of
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Society. Institutions may subscribe to the Journal but may not become members.
Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests.
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special interests. There are four numbers in each volume of the Journal, scheduled
for February, May, August and November, and eight numbers of the News each year.

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Second class postage paid at Lawrence, Kansas, U.S.A. 66044.

fag

JOURNAL OF

Tue LeprpopreristTs’ SOCIETY

Volume 25 1971 Number 2

A REVIEW OF THE GENUS ARCAS WITH DESCRIPTIONS
OF NEW SPECIES (LYCAENIDAE, STRYMONINI)

S. S. NICOLAY
1500 Wakefield Drive, Virginia Beach, Virginia

In 1832, William Swainson described Arcas as a “subgenus” of the sub-
family “Theclanae” (=Theclinae) and established Papilio imperialis
Cramer as the type species. Yet, most writers throughout the intervening
138 years have neglected to use the name, to identify additional species
as belonging to this subgenus or to further define and clarify its status.
The purpose of this work is to establish Arcas as a valid genus of the
Lycaenidae, and to separate those species that belong to Arcas from the
all-inclusive genus Thecla.

Arcas, as presently conceived, contains seven species; five heretofore
contained in Thecla Fabricius, the sixth and seventh described as new.
The male of one species, A. splendor Druce has been unknown for almost
64 years and is herein described and figured for the first time. Clench
(1963) allied Arcas with Atlides Hiibner and Pseudolycaena Wallen-
gren. Certainly the three genera share many structural similarities. By
the same token, Arcas may be readily separated from both by such sig-
nificant characters as the male scent-spot, the very deep tornal cleft and
extraordinarily long tails on the hind wing, and obvious differences in the
genitalia of both sexes. All species in the genus are Neotropical. The type
species, A. imperialis has the widest distribution, being found virtually
unchanged in habitus from Mexico south through Panama and throughout
South America to southem Brazil and Bolivia (Figure 6).

The following abbreviations are used to indicate the collections from
which specimens have been examined and data recorded in this study:
(USNM) Smithsonian Institution, Washington, D.C.; (CM) Camegie
Museum, Pittsburgh, Pennsylvania; (AM) American Museum of Natural
History, New York; (AF) Allyn Museum of Entomology, Sarasota, Flor-
ida; (MN) Museu Nacional, Rio de Janeiro, Brazil; (GS) Gordon B.

88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Small collection, Balboa, Canal Zone; (PG) P. Gagarin collection in Rio
de Janeiro, Brazil; (JD) Dr. John C. Downey collection, Cedar Falls,
Iowa; (N) the author's collection.

Genus Arcas Swainson, 1832
Type species: Papilio imperialis Cramer, 1775

Hind wing with two long tails, the shorter (5 mm) at the end of Cui, the second
at the end of Cuz at least twice as long (10-15 mm); anal angle very deeply cleft
forming an elongate, narrow anal lobe. Male with a scent-spot on forewing. Ab-
domen yellow beneath. Palpi very long, thick, porrect and twice as long as head,
curved downward, with all joints entirely covered with close-set mixed blue and
black scales. Frons and head covered with brilliant green metallic scales; antennae
black, slender, the club scarcely thicker than stalk; eyes with short, obscure bristles.

Male genitalia with the saccus very long, slender, almost twice as long as complete
genital ring; ventrally, a pair of pointed, triangular processes extending from the an-
terior vinculum, and curving partially around valvae; valvae rather compact, broad,
about one-half length of vinculum, separated their entire length. Aedeagus very
long, longer than combined length of vinculum and saccus, slender, apically flaired
and upturned, with two terminal cornuti, one with obscure terminal apical teeth.

Female genitalia with a complex ductus bursae consisting of three distinct sections
or elements; entire terminal section of ductus bursae cone-shaped, dorsal plate
heavily sclerotized, fan-shaped, relatively wide, smooth-edged, ventral portion mem-
branous near opening, becoming sclerotized at base; mid-section a long, curved, very
lightly sclerotized narrow tube; anterior section a sclerotized, widening, recurved
complex structure providing entry into corpus bursae; latter a large, rounded sac,
adorned by two simple, single-spined signa on inner surface.

As is the case in many Strymonini genera, the genitalia of all species
in Arcas are very similar. Furthermore, the genitalia of both sexes show
considerable individual variation within a single species. It would indeed
be difficult to separate species in this genus on the basis of the genitalia
alone. A ventral view of the full male genitalia with the aedeagus re-
moved is shown in Figure 2, and a lateral view with aedeagus in place is
illustrated in Figure 1. With the exception of the valvae, illustrated for
each species, no other consistently recognizable differences between
species could be found in the male genitalia of the genus. The female
genitalia also show considerable individual variation within each species,
and a striking similarity among the seven species. Illustrated are ventral
views of the complete genital complex for the two new species, a lateral
view of A. ducalis and a ventral view of the terminal cone-shaped section
of the ductus bursae of the remaining species.

The male genitalia of Arcas species, although somewhat similar to those
of the genus Atlides, are consistently different in that the latter are more
stoutly built, the saccus shorter, broader, the aedeagus heavier, the valvae
very much broader, and in some species, joined together near the anterior
end, The female genitalia of species in the two genera show little similar-

VOLUME 25, NUMBER 2 89

Fig. 1. Male genital structures of Arcas Swainson. a, Lateral view of complete
genitalia of A. jivaro Nicolay; b, ventral view of valvae and lateral view of terminal
end of aedeagus of A. imperialis Cramer; c, same views and structures of A. ducalis
Westwood; d, same views and structures of A. cypria Geyer.

90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 2. Male genital structures of Arcas Swainson. a, Ventral view of genitalia
with aedeagus removed of A. delphia Nicolay; b, lateral view of complete aedeagus
of A. delphia Nicolay; c, ventral view of valvae and lateral view of terminal end of
aedeagus of A. tuneta Hewitson; d, same view and structures of A. splendor Druce.

VOLUME 25, NuMBER 2 91

Fig. 3. Female genital structures of Arcas Swainson. a, Lateral view of complete
genitalia of A. ducalis Westwood; b, ventral view of above genitalia, showing terminal,
cone-shaped portion of ostium bursae with its fan-shaped dorsal plate; c, ventral view
of the terminal portion of the ostium bursae of A. imperialis Cramer.

ity. In contrast to the smooth-edged dorsal plate and long, curved, com-
plex ductus bursae in Arcas, species in Atlides have a ductus bursae with
a spined dorsal plate and a heavily sclerotized, simple straight tube en-
tering directly into the bursa copulatrix. The two genera are similar in
that both have a pair of single-spined signa on the interior surface of the
bursa copulatrix.

The male genitalia of Arcas show consistent and obvious differences
from those of Pseudolycaena in the size, shape and structure of the uncus,
saccus and valvae. In Pseudolycaena, two heavily sclerotized, toothed
processes extend from the ventral surface of the valvae but are completely
lacking in Arcas. Female genitalia in the two genera differ in many re-
spects, but primarily in the shape and length of the tubular ductus bursae

92 JouRNAL OF THE LEPIDOPTERISTS’ SOCIETY

Fig. 4. Female genital structures of Arcas Swainson. a, Ventral view of complete
genitalia of A. delphia Nicolay; b, ventral view of terminal section of ostium bursae of

A. tuneta Hewitson; c, ventral view of the terminal section of ostium bursae of A.
cypria Geyer.

which, in Pseudolycaena is heavily sclerotized, simple, and relatively
straight, emptying directly into the bursa copulatrix, in contrast to the
very complex, curved and recurved structure in Arcas. Both genera have
a bursa copulatrix adorned with two single-spined signa.

Arcas contains some of the most lavishly formed and strikingly colored
butterflies in the Neotropics. Eagerly hunted by most collectors, many of
the species are still not common in collections. Generally, they are “hill-
toppers, in that the males can be found at the summit of the highest point
of land within a given area. This trait, combined with a preference for a
lofty tree perch makes for a difficult catch at best. Nothing is known of
their life history. Some species are true rarities. All collections I have

VOLUME 25, NuMBER 2 93

Fig. 5. Female genital structures of Arcas Swainson. a, Ventral view of complete
genitalia of A. jivaro Nicolay; b, ventral view of terminal section of ostium bursae of
A. splendor Druce.

studied contained relatively few specimens of A. tuneta and A. delphia,
with the latter always included within the series of tuneta. The type and
allotype of A. jivaro are the only specimens known. A. splendor remained
“uncollected” for well over 100 years until finally discovered again by

94 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Cee

Fig. 6. Distribution of species of Arcas. Solid circles, A. imperialis Cramer; open
circles, A. cypria Geyer; half-solid circles, A. delphia Nicolay; up-right solid triangles,
A. ducalis Westwood; inverted solid triangles, A. splendor Druce; open circle with
slanted line, A. tuneta Hewitson; inverted solid triangle on line, A. jivaro Nicolay.

Gordon Small in 1964. The species is still known from only two rather
isolated localities in Central America.

Key to the Species of Arcas

1 Underside hind wing without a median line
Underside hind wing with a median line 3

Hind wing underside brilliant metallic green, striated with black (females
from Colombia, Panama and Central America perhaps with a pink wash on
Miscye eS Te ie a A. imperialis (Cramer )

Hind wing underside brilliant metallic green, with a clearly defined carmine
disc and wide yellow-gold outer wing margin A. ducalis (Westwood )

Upperside of forewing, male with scent-spot tiny, located well outside the cell,
not touching transverse vein; female upperside with dark margins of both
wings very ‘wide’ 2s) Fe) Pe eee A. delphia Nicolay

i)

Ww

VoLUME 25, NUMBER 2 95

Upperside of forewing, male with scent-spot touching or bisected by the trans-

PEESS VSD cee ae ee ee eee ere ee. Sane eee te Re eaneeeie ar 4

4 Underside of hind wing with median line bent sharply distad at vein Ms: before
curving irregularly to the inner margin _...____..---__----- A. cypria (Geyer)

Underside of hind wing with median band not bent distad, but straight or
MehenyZCOncave tO interspace Cus 5

5 Upperside of forewing blue, male with scent-spot large, crossing the transverse
eIUURE OME WC 1CC limemernemnie men sas fe ge 6

Upperside of forewing green, male with scent-spot small, lying outside the cell,
touching the transverse vein; female with dark outer wing margins rela-

CSE PSLS STITT ONNY cae a ele A. tuneta (Hewitson )
6 Upperside of forewing, male with scent-spot sharply defined, tear-drop shaped
with the proximal point inside the cell; female with underside of forewing
with a dark post-median line; the largest species in the genus ________--_______-

ce cnn ER A. splendor (Druce)
Upperside of forewing, male with scent-spot filling distal half of cell extending
beyond transverse vein, vaguely defined with a greenish halo; female with

no post-median line on underside of forewing ___........--____-. A. jivaro Nicolay

Arcas imperialis (Cramer )
Figs. lb, 3c, 7A, 7B, 10A.

Papilio imperialis Cramer, 1775, p. 120; pl. 76, figs. E, F.

Papilio venus, Fabricius, 1781, p. 115.

Theritas venus, Hiibner, 1819, p. 80.

Arcas imperialis, Swainson, 1832, p. 88.

Eucharia imperialis, Boisduval, 1870, p. 14.

Thecla imperialis, Hewitson, 1877; 1: 71. Godman & Salvin, 1888; 2:13, 3; tab. 48,
figs. 15 and 16. Draudt in Seitz, 1921; 5: 748, pl. 146c. Comstock & Huntington,
1960; 68: 234.

Thecla oakesii, Butler, 1884; 14: 267.

“Above shining blue: beneath emerald-green, marked with minute black waved
lines. . . It is impossible to depicture with correctness, the resplendant blue which
ornaments the upper surface, or the vivid emerald green on the underwings, of this
rare and splendid insect.”

Little need be added to the above quotation from Swainson’s remarks
as he described the type species, Cramer’s P. imperialis. It is a favorite
species for color illustrations, and would be difficult to confuse with any
other lepidopterous insect.

The black apical border on the upper forewing is large and is more
sharply defined in imperialis than in other species in the genus. The dark,
red-brown scent-spot on the male forewing is placed at the cell end, half
in and half outside the cell, the transverse vein clearly marked by a line
of brilliant blue scales ( Fig. 10A ).

Imperialis has a wide geographical distribution from Mexico southward
through Panama and through South America to southem Brazil and
Bolivia.

96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

| Fig. 7. Arcas spp. A, Arcas imperialis (Cramer) 6, Rio Bodoquero (1300 ft.)
Caqueta, Colombia, 19 Jan. 1969 (S. S. Nicolay); B, underside of specimen in A;
C, A. ducalis (Westwood), 6, Joinville, Santa Catarina, Brazil, 28 Oct. 1967 (O.
Mielke); D, underside of specimen in C; E, A. cypria (Geyer), 6, Los Rios, Canal
Zone, Rep. of Panama, 29 Jan. 1965 (S. S. Nicolay); F, underside of specimen in E.

VOLUME 25, NUMBER 2 7

Specimens Examined. Mexico: Vera Cruz—Presidio (USNM, AM) 292. Guate-
mala: Cayuga (CM) 19. Honduras: No locality (USNM) 192. Costa Riea:
Guapiles (USNM) 16. Carillo (USNM) 1¢. Panama: Bugaba (USNM) 1é4 19.
Canal Zone—Los Rios, Madden Forest, Summit, Barro Colorado Isl. (AM, GS, N)
10$ 72. Colombia: Cundinimarca—Bogota (USNM, CM) 2¢é. Magdalena—Minca
(CM) 168. Antioqua—Casabe (AF, AM) 39, Rio Cocorna (AM) 12, Medellin (AF)
26 12. Caqueta—Rio Bodoquero; Tres Esquinas (N) 2¢. Amazonas—Rio Tacana
(AM) 16. Vague—Felip Ovalle (AM) 13¢ 89. El Centro, Magelena Valley
(AM) 2¢. Melgar (CM) 18. Yacopi (USNM) 19. No locality (USNM, AM, MN)
4¢ 19. Venezuela: Las Quiguas, Esteban Valley (CM) 4¢. Cucuta (USNM) 19°.
Peru: Loreto—Iquitos, Putumayo River, Pucallpa, Achinamiza (AM, AF) 16¢ 19.
San Martin—BellaVista, Tarapoto (AM, PG) 24. Cusco—La Salud (CM) 292.
Amazonas—Rio Santiago (AM) 12. Bolivia: Las Juntas, Rio Surutu, Rio Yapacavi,
Prov. del Sara, Buenavista, Cuatro Ojos (CM) 14¢ 29; Rurrenebaque 14 19; San
Pedro (AM) 1. French Guiana: Pied Saut., Oyapok River, No locality (CM) 2°.
Brazil: Pard—Santarem, Obidos, Utinga (Belem) (AF, MN) 3é 22. Amazonas—
Ipiranga, Rio Maues, Manacopuru, Teffe, Rio Solimoes, S$. Paulo de Olivenca, Juarete,
Rio Negro (CM, MN, AM, N) 136 42. Mato Grosso—Buriti, Chap. Guimaraes (N)
1g. Minas Gerais—Paracatu (N) 2¢. Goias—Jatai, S. Rita Araguaia (N) 2é4.
District Federal—Parque do Gama (N) 22. Espiritu Santo—No locality (MN) 1¢.
Rio de Janeiro—Independencia, Petropolis, Nilopolis, Araruana, Angra dos Reis,
Boca do Mato (MN, PG) 5¢ 32. Guanabara—Gawea, Paineiras, Corcovado, Rio de
Janeiro (PG, MN, AF, USNM) 64 32. Sao Paulo—Mendes, Loreto (MN, AF) 2¢.
Parané—Caviuna (AM) 1°. Santa Catarina—Massaranduba-Blumenau, Annaburg,
Joinville (MN, USNM, AM, N) 10¢ 42. Not located: Colana Island (USNM)
KE=SNodata:- 146 29.

I have taken this beautiful insect in Panama, Colombia and Brazil. Its
flight habits and specific haunts are similar in each locality in which I
have found it. The male sits perched on a leaf 15-20 feet high above a
small sunlight clearing in the heavy forest, whence it makes infrequent
and short, swift flights, to return to the same spot. Both sexes may occa-
sionally be found on flowers and, early in the mornings, females at the
sunlight edge of the heavy forest on low bushes. Past authors have called
it “rare.” It is not a common insect, but I would suggest its rarity is due
in large measure to its flight habits, for it is difficult to locate and capture.

The name Thecla oakesii Butler was given to specimens with a coppery
or rosy wash on the disc of the hind wing beneath. Its position in past
literature is confusing and variable. Draudt (1921) made it a synonym
of ducalis Westwood. Lathy (1930) made it a female form of the “Colom-
bian race” of imperialis. T. oakesii is not a synonym of ducalis and I am
not at all sure that Lathy’s treatment is entirely correct. I have noted that
some female imperialis taken in localities other than Colombia have vary-
ing degrees of the rosy wash on the disc of the hind wing beneath. Of
the series of six specimens labeled T. oakesii in the National Museum col-
lection, 4 are females, one each from Mexico, Panama, Venezuela, and
Colombia. The 2 males, both from Colombia, have a very small and only
the faintest indication of pink tint in the disc. The substantial series of

98 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

males from Colombia in the American Museum show no pink wash what-
soever; nor do any of numerous specimens I have seen from Panama show
this pink wash in the disc. It appears that the amount of rosy tint in the
disc on the underside of the hind wing is a highly variable characteristic
found primarily in female imperialis taken in Colombia, Venezuela and
northward into Panama and Central America. The name oakesii Butler
is not needed.

Arcas ducalis (Westwood )
Figs. 1c, 3a, 3b, 7C, 7D; 10B.

Thecla ducalis Westwood, 1852; 2: 483, pl. 77, fig. 1 (figure only). Kirby, 1879, p.
151. Draudt in Seitz, 1921; 5: 746, pl. 146c. Lathy, 1930; 78: 133, pl. 9, figs. 3
and 4. Comstock & Huntington, 1959; 67: 211.

The original figure of this species was published without an accom-
panying written description. In 1930, Percy I. Lathy wrote concerning
the relationship of ducalis:

“There has been considerable confusion in collections respecting this species. It
is one of those cases where a species has been figured but there is no accompanying
description, and as only the upper side is shown the figure might apply to the female
of imperialis Cram. As a matter of fact Druce took the female of oakesii, Btl., a
local race of imperialis to be ducalis. Draudt in Seitz, p. 746, also regards ducalis
as a race of imperialis and places oakesii as a synonym. This is incorrect as a glance
at the figures . . . will show. T. ducalis is a quite distinct species and not a race of
imperialis, as the two occur together in South Brazil. . .”

The brilliant metallic carmine disc and wide yellow-gold outer margin
on the underside of the hind wing are the most obvious macular differences
between ducalis and imperialis. Yet, there are additional and consistent
differences between the two. Although the forewing apex in imperialis
is rounded, it never approaches the extreme foreshortened appearance of
ducalis; the forewing of ducalis is almost elliptical. The male scent-spot
in imperialis is bisected by narrow blue scaling clearly visible on the
transverse vein of the cell end; in ducalis the primary scent-spot is un-
broken, lying outside the cell, with a tiny group of pale brown scales of
a different texture grouped within the cell (Fig. 10B). In imperialis the
upper hind wing terminal margin is a well defined thin black line, slightly

wider at the wing apex; in ducalis this margin is vaguely defined and
wider throughout the entire wing margin. And although it is a subtle
difference, more obvious in males than females, the basic upper side wing
color in ducalis is a deep blue, without the greenish hues of imperialis.

The geographical distribution of ducalis is restricted to the states of
southern Brazil. At some svecific localities within these states, ducalis
and imperialis have both been taken e.g. Independencia near Petropolis,
Rio de Janeiro and at Joinville and Jmvarana.

VOLUME 25, NuMBER 2 99

Specimens Examined. Brazil: Rio de Janeiro—Teresopolis; Independencia,
Petropolis (MN, PG, USNM, N) 4é¢ 22. Sao Paulo—Umvarana (PG) 19. Parana
—Castro; Londrina; Caviuna (AM, USNM) 32 192. Santa Catarina—Neudorf;
Nova Teutonia; Joinville; Massaranduba-Blumenau (MN, AM, PG, N) 46 89.
Rio Grande do Sul—No locality (MN, AM) 246 19.

Arcas cypria (Geyer )
lmies, lel, Ze, 71a, We, ICID).

Theritas cypria Geyer, 1837 in Hubner, 1837; 5: 36, figs. 945 and 946.

Pseudolycaena paphia, Felder, 1864-1867; 2: 234, pl. 28, figs. 12 and 13.

Thecla cypria, Hewitson, 1877; 2: 71. Godman and Salvin, 1887-1901; 2: 13, tab.
XLVIII, figs. 12 and 13. Draudt in Seitz, 1921; 5: 746, pl. 146d. Comstock &
Huntington, 1959; 72: 201.

Specimens of this species have been carefully compared with the orig-
inal description and with the colored plates that accompanied it. The
colored figures by Godman and Salvin (1887) are very well done, but the
Seitz (1921) figure of the male upperside lacks the scent-spot on the fore-
wing, and the entire figure is done in green rather than the basic blue
color of this species on the upper surface. The male dark brown scent-spot
is located within the cell with a blue-grey patch of scales of a different
texture adjoining outside the cell across the transverse vein (Fig. 10D).
The black-brown margin is wide and vaguely defined on both upper wing
surfaces.

On the hind wing beneath, a wide black median line, proximally etched
in glossy white, begins about mid-costa, is bent distally at vein Ms, then
is curved irregularly in an arc to the midpoint of the inner margin. From
the median line to the base, the black-flecked brilliant green scaling is
washed with a shining coppery-gold. The apices of both fore- and hind
wings are dusted in glossy white. The remainder of the hind wing is
brilliant green, irregularly dusted with black scaling.

Felder (1867) gave the name paphia to specimens of this species with
a coppery red disc on the underside of the hind wing. Godman and Sal-
vin (1887) placed paphia as a synonym of cypria. I agree with this treat-
ment of the name. The series of cypria in the National Museum and
American Museum collections were taken in a variety of locales from
Mexico to Panama. It would be difficult indeed to separate out and des-
ignate a subspecies from this group on the basis of a coppery wash on
the disc of the underside of the hind wing. Gordon Small and I have both
taken this species in various localities in Panama; the intensity and degree
of coppery wash seem to vary with individual specimens rather than being
the mark of a different species or even a subspecies.

Cypria is essentially a Central American species, ranging from Mexico
south to Colombia.

100 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Specimens Examined. Mexico: San Luis Potosi—Ciudad Valles (CM) 19.
Vera Cruz—Paos San Juan; Coatepec; Presidio; Jalapa; Cordoba City; Misantla
(USNM, AM) 88 59. Chiapa—Tapachula; Palenque (AF, CM, AM) 44.
Tabasco—Terrosique (AM) 34. Yucatan—Piste 15; Chichen Itza (CM) "Sige Do.
Quintana Roo—Xcanha (CM, AM) 24. Guatemala: Cayuga (USNM, CMO?
Sayaaxche, El Peten (AM) 26 19. Costa Rica: Port Limon (USNM) 1¢. Tur-
rialba, Cartago Prov. (JD) 1é. Panama: Chiriqui (USNM, AM) 24. Arraijan
(AM, N) 24. Cerro Campana (GS, JD, N) 36 49. Canal Zone—Los Rios;
Madden Forest; Cocoli (GS, N) 11é 19. Colombia: Medellin (AF) 16. El
Centro, Magdalena Valley (AM) 246 29.

Arcas tuneta (Hewitson)
Figs. 2c, 4b, 8E, 8F, 10E.
Thecla tuneta Hewitson, 1865; 1: 72, vol. 2, pl. 28, figs. 14 and 15. Draudt in Seitz,

1921; 5: 746, pl. 146d. Comstock & Huntington, 1961; 72: 177.

The Hewitson (1865) figures are excellent, and the Seitz (1921) illus-
trations are a reasonable likeness of a female. The upper wing surface of
the male is more greenish than any other species in the genus. The black-
brown outer margin is only modestly wide on the forewing, beginning at
the middle of the costal margin, thence curved out beyond the cell to the
tornus and includes the apex and all of the outer margin. The scent-spot
is small, inconspicuous, and lies outside the cell, its inner margin touching
the transverse vein (Fig. lOE). There is no secondary or adjacent patch
of androconial scales. The hind wing margin is moderately broad, black-
brown and vaguely defined, and reaches from the apex to the first tail at
vein Cu;. Two black-brown spots lie at the base of the tails.

The underside of the hind wing is dark metallic green, heavily irrorate
with black scaling distad to the median line, the inner or proximal half
slightly touched with gold scaling. The median line is black and runs
almost straight and unbroken from the costal margin to vein Cus, then
is bent sharply in a short line to the mid-point of the inner margin.
Although the median line is slightly concave in tuneta and uneven along
the proximal margin, it does not break distally at any point as it does in
cypria.

Tuneta is apparently one of the more uncommon species of this genus.
Hewitson states merely that the species is from South America. Near
Petropolis, Rio de Janeiro it flies at Independencia where imperialis and
ducalis have also been taken. Bahia is the most northern locality in Brazil
whence tuneta has thus far been recorded. Dr. Keith Brown took it here
on 15 Dec. 1966 at Ubata on the Rio Jequie in a section of deep forest near

a hilltop at 300 meters, a location on the rain border between Amazon
coastal forest and dry caatinga.

Specimens Examined. Peru: San M artin—Juanjui; Yumbatos (AM) 16 19.
Putumayo River (AM) 19. Bolivia: 0 Surutu; Cuatro Ojos; Portachuelo, Rio

VOLUME 25, NUMBER 2 101

Palmetillas (CM) 36 29. Brazil: Bahia—Ubata, Rio Jequie (KB) 1é. Rio de
Janeiro—Independencia, Petropolis (PG) 4¢ 12. Sao Paulo—No locality (USNM)
12. Santa Catarina—Joinville; Massaranduba-Blumenau (PG, GS, AM, N) 4é¢ 29.

Arcas delphia Nicolay, new species
Figs. 2a, 2b, 4a, 8A, 8B, 8C, 8D, 10F.

Male. Length of forewing, 19 mm. Upperside: Basic wing color brilliant blue-
green; forewing margin black-brown, beginning inside midpoint of costal margin,
thence curved outside scent spot to tornus, including all of apex and outer margin;
scent-spot very tiny, lying completely outside cell, well beyond transverse vein.
Hind wing brilliant blue-green with outer margin a thin black line; apex and costal
margin vaguely defined, broader, black-brown; a single dark anal spot in interspace
Cuz. Underside: Forewing dark forest green, irrorate with sparse black scales, disc
pale blue from vein Cuz to inner margin; outer margin a thin black line, fringes nar-
rowly pale blue. Hind wing dark, brilliant forest green dusted with sparse black
scaling, more heavily so distad of the median line. Median line heavy, black, beginning
at costal margin, running almost straight to interspace Cuz, thence bent sharply at a
90° angle straight to inner margin; outer margin a thin black terminal line from costa
to anal lobe. Anal Jobe and tails black; fringes pale blue.

Female. Length of forewing, 20 mm. Upperside: Forewing blue with a scattering of
green scaling, the margin very wide, black-brown running narrowly from base along
costa, thence curving around end of cell to tornus. Hind wing blue with a wide, vaguely
defined black-brown margin to vein Cui; a dark spot in each of interspaces Cui and
Cuz. Anal lobe and adjacent interspace Cuz metallic green; fringes black. Underside:
Forewing dark green irrorate with sparse black intermixed yellow-gold scales; space
from vein Cuz to inner margin grey-brown; fringes and terminal line, black. Hind wing
dark green, with intermixed gold scaling in disc and heavily dusted with black be-
tween median line and outer margin; median line slightly concave, bent sharply to in-
ner margin at vein Cuz; a thin black terminal line running from apex to anal lobe; anal
lobe black; fringes narrowly pale blue.

Holotype, male, Guapiles, Prov. Limon, Costa Rica, 850 ft., August (year and col-
lector unknown). Allotype, female, Finca la Lola, vic. Madre de Dios, Limon Prov-
ince, Costa Rica, 21 July 1965. Paratypes in the Smithsonian Institution collection:
one male, Guapiles, Costa Rica, 850 ft., one male, Yacofsi, Colombia; in the American
Museum of Natural History: one female, Costa Rica (no additional data). A single
male and three female paratypes in Mr. Gordon Small’s collection from the allotype
locality were collected on 20-23 July 1965 and a female from Gamboa, C. Z., 25 June
1970. In Dr. Downey’s collection are one male and two female paratypes taken at
the allotype locality on 19 July 1965; in the author’s collection, a single male para-
type, Colon (Santa Rita), 1500 ft., Panama, 2 February 1969 and three female para-
types from the allotype locality in Costa Rica. In the Schmidt-Mumm collection in
Bogota, Colombia is a male taken at Victoria, Dept. of Caldas in August 1958.
The male holotype is deposited in the Smithsonian Institution, Washington, D.C., the
female allotype, in Mr. Gordon Small’s collection, Balboa, Canal Zone.

It is obvious that delphia is closely related to tuneta. The genitalia
of both sexes are very similar with no apparent recurring or obvious dif-
ferences. Yet, the location of the male scent-spot, longer hind wing and
much reduced dark margins on the upper hind wing of the male delphia
remain consistent differences between the two species. Females are more
difficult to separate, the most consistent difference being the very wide

JouRNAL OF THE LEPIDOPTERISTS SOCIETY

bo

10

Mig. 8. Arcas spp. A, Arcas delphia Nicolay, ¢, paratype, Finca la Lola, vic.
Madre de Dios, Limon Proy., Costa Rica, 20 July 1965 (G. B. Small); B, rind exon:
of specimen in A; C, A. delphia Nicolay, 2, allotype, same locality as paratype in
\, 21 July 1965 (G. B. Small); D, underside of specimen in C; E, A. tuneta (Hewit-
son), 2, Joinville, Santa Catarina, Brazil, 28 Oct. 1967 (O. Mielke); F, underside of

pecimen in

VoLUME 25, NuMBER 2 103

Fig. 9. Arcas sp. A, Arcas splendor (Druce), 6, Cerro Campana (2500 ft.),
Rep. of Panama, 5 Aug. 1964 (G. B. Small); B, underside of specimen in A; C, A.
splendor (Druce), @,; Cerro Campana, Rep. of Panama (2500 ft.) 29 Aug. 1966 (G.
B. Small); D, underside of specimen in C.

heavy dark margins on the upper surface of both fore- and hind wings in
delphia.

Like cypria, delphia is essentially a Central American species, but ob-
viously quite rare. It has been found no further north that Costa Rica
with its southern limits in Colombia.

Arcas splendor (Druce)
Figs. 2d, 5b, 9A, 9B, 9C, 9D, 10C.
Thecla splendor Druce, 1907, p. 570; pl. 31, fig. 4. Draudt in Seitz, 1921; 5: 746,
pl. 146d. Comstock & Huntington, 1961; 71: 196.
For over 100 years, splendor has been known and represented in col-
lections by only Druce’s type, a worn female, taken some time between
1848 and 1857. In the summer of 1964, Mr. Gordon Small took a series of

104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

D

Fig. 10. Upper right forewing of male of Arcas spp. showing scent-spot. A, Arcas
imperialis (Cramer) 8; B, A. ducalis (Westwood) 6; C, A. splendor (Druce) ¢;
D, A. cypria (Geyer) 6; E, A. tuneta (Hewitson) ¢; F, A. delphia Nicolay ¢.

males and females of this species in the Republic of Panama, thus record-
ing for the first time, the capture of a male of this splendid insect. Druce's
original description is of interest and quoted herewith:

“Female. Allied to T. tuneta Hew. which it closely resembles on the upper side.
On the under side it differs in possessing a curved, black, ultramedian band on the
fore wing, inwardly edged with whitish, and in the median band on the hind wing
being broader, placed further from the base, more concave, and more sharply angled

to the anal margin. There is also a submarginal indistinct dark shade which is not
present in T. tuneta Hew.

“Expanse 1%o9 inch.

“Hab. Colombia

“Type, Mus Oxford, No. 1901/1198

“Collected 1848-1857 by H. M. Vice-Consul Edward W. Mark, and presented to
the Museum in 1901 by Mr. F. W. Mark.

“The type, which is the only specimen I have seen, is not in very good condition,
having lost its antennae and abdomen; but enough remains to show that it is a very
distinct species, and I believe unnamed.”

Male. Length of forewing, 22 mm +2 mm. Upperside: Forewing brilliant blue
with golden-green scaling intermixed; the black-brown margin moderately wide with a
vaguely defined inner edge, beginning about mid-costa then curving outside scent-spot
to tornus and including all of apex and outer margin; scent-spot tear-drop shaped, large,
black brown, placed with proximal point inside cell, remainder of spot outside cell
(fig. LOC), Hind wing of same brilliant blue-green color with a black margin wider
at apex, narrowed to a thin black line at tornus, with black spots at tornus in inter-

paces Cu, and Cu. Underside: Forewing dark green with scattered black scales, a

' .
narvmna

| line and pale blue fringes. Hind wing the same deep green; a
ecdian line, slightly concave with a thin white proximal margin, running

VoLUME 25, NUMBER 2 105

from costal margin to mid interspace Cuz, then sharply bent at 90° angle straight to
inner margin. A thin black terminal line beginning at apex and ending at black anal
lobe; fringes pale blue. Area between median line and terminal line, heavily smudged

and dusted with black.

Draudt (1921) presumed splendor to be allied to tuneta, referring to it
as “probably only a form...” I have examined the type in the British
Museum, comparing it carefully with color transparencies of specimens
collected by Gordon Small in Panama. I have no doubt that the Panama
specimens are A. splendor and that splendor is indeed a valid species.
The female of splendor is unique in the genus, in that it has a post median
line on the underside of the forewing. The male is easily separated from
those of other species in the genus by the large tear-drop shaped scent-
spot on the forewing. Splendor is the largest species in the genus.

It would be impossible at this time to pinpoint the type locality more
accurately than “Colombia.” Yet, it is worthy of note that Panama was a
part of Colombia when Druce’s type was collected. The currently known
geographic range of splendor is limited to Panama and Costa Rica. Here,
limited to two specific localities, Cerro Campana (2500 ft.) Panama and
Moravia (3500 ft.) Cartago Prov., Costa Rica, it flies in the rain forest
that clothes the mountain tops. We have searched but have not been able
to find this species during recent collecting trips to many other localities
in both Colombia and Panama.

Specimens Examined. Panama: Cerro Campana, 2500 ft., Panama Prov. (GS, JD,
N) 193 59. Costa Riea: Moravia, 3500 ft., Catago Prov. (GS, JD) 1¢6 29.

Arcas jivaro Nicolay, new species
Bigseeias od evAS LIB biG: hb:

Male. Length of forewing, 17 mm. Upperside: Basic wing color brilliant blue with
a slight greenish cast; forewing dark margin wide, with a well defined inner margin
beginning just proximal to midpoint of costa, curved outside cell and scent-spot and
then to tornus, including all of the apex and outer margin; the scent-spot large, but
vaguely defined, filling distal half of cell, extending out beyond transverse vein and
surrounded by a greenish halo. Hind wing the same brilliant blue, with a very narrow
dark outer margin, vaguely broader near apex and along costa; a dark narrow bar
across anal angle at interspaces Cuz and Cu:. Underside: Forewing dark lustrous
green with pale-blue scaling in disc above inner margin; outer margin a thin black
terminal line, fringes dark grey-blue. Hind wing dark lustrous green, sparsely irrorate
with black, most prevalent distad of median line toward anal angle. Median line
black, proximally etched with a very thin line of light scales, extending from midpoint
of costa almost to cleft of anal lobe, then sharply bent at 90° angle to inner margin.
Outer margin a thin black terminal line from vein Rs to anal lobe; anal lobe and tails
black, fringes narrowly pale blue.

Female. Length of forewing, 18 mm. Upperside: Forewing lustrous blue with a
vaguely defined, wide, dark margin beginning just proximal to mid-point of costa
then curved around outside cell to tornus. Hind wing lustrous blue with a vaguely
defined, dark outer margin, widest just below apex, disappearing at Cu; terminal
margin a thin black line from apex to anal lobe; anal lobe and adjacent interspace

106 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 11. Arcas sp. A, Arcas jivaro Nicolay, ¢, holotype, Sucula, Macas, Ecua-
dor (800 m) date and collector unknown; B, underside of specimen in A; C, A. jivaro
Nicolay, 2, allotype, same data as holotype; D, underside of specimen in C.

iridescent green. A dark narrow bar across interspace Cus, a dark spot in interspace
Cu, fringes narrowly blue. Underside: Forewing lustrous golden-green, disc above
inner margin light brown out to and including tornus; terminal line black, thin; fringes
brown. Hind wing golden green in disc proximal to median line; dark green heavily
dusted and smudged with black just distad of median line toward anal angle. Median
line black, etched proximally in white, extending from mid-costa to just above cleft
of anal lobe, then curved sharply at 90° angle to inner margin. Anal lobe and tails
black; terminal line black, narrowly etched in white from above costa to Cu, extend-
ing from costa to anai lobe; fringes very pale blue, almost white.

Holotype, male, Sucula, Macas, Ecuador, 800 meters (date and collector un-

known). Allotype, female, same data. The holotype and allotype are located in the
American Museum of Natural History, New York.

Some interesting observations can be made about this species. Among
the many dozens of specimens of Arcas I have studied, representing most
of the major collections in ‘his hemisphere, these are the only two speci-
mens of the genus I have seen from Ecuador. Primarily I presume, this

VoLuME 25, NuMBER 2 OM

is due to the lack of collecting and/or collectors in Ecuador. Yet, even
imperialis is notably lacking in collections. The male of A. jivaro displays
a rather striking mixture of characteristics found in other species of the
genus. The underside of both sexes, but particularly that of the female is
similar to tuneta. Yet, the upperside of the male is the basic lustrous blue-
green of imperialis, not the green of tuneta. The wide, rather sharply de-
lineated dark wing margins of the forewing are also like those of im-
perialis. The underside of the hind wing is the dark lustrous green of A.
splendor with a relatively narrow but heavy black median line, also rem-
iniscent of that species. The scent-spot is unlike that of any other species
in the genus. The single female is, however, indistinguishable from fe-
males of tuneta. It is possible that a small series of females of jivaro
would reveal some consistent and reliable differences.

The presently known geographic distribution of A. jivaro is restricted to
the type locality, located near the town of Macas, the capital of the Ter-
ritorial Division of Morona Santiago, Ecuador.

Acknowledgments

I wish to thank a number of people who provided the information and
valuable assistance that is so vital to a work of this scope. To Gordon B.
Small, Jr., Balboa, Canal Zone and Dr. John C. Downey, University of
Northern Iowa, Cedar Falls, Iowa must go credit for the excellent field
work which provided the basis for much of the new information con-
tained herein. I am indebted to Dr. Keith S. Brown, Jr., Rio de Janeiro,
Brazil who provided a great deal of the distributional data and other help-
ful information on the material in the Museu Nacional collection.

Mr. T. G. Howarth and Mr. G. E. Tite generously allowed me to
examine types in the British Museum. William D. Field, United States
National Museum; Dr. F. H. Rindge, American Museum of Natural His-
tory; Harry K. Clench, Carnegie Museum and Dr. Lee D. Miller, Allyn
Museum of Entomology, provided the assistance and cooperation that al-
lowed me to examine the material in the collections in their care, and of-
fered many helpful suggestions during the course of this study.

The photographs were made by Robert C. Williams, formerly of the
U.S. Marine Corps, now at the University of Michigan and Lt. G. G.
Thomas, USMC. Their interest, energy and technical competence were
especially helpful. Drawings of the genitalia were made by the author.

Literature Cited

BoispuvAL, JEAN A. 1870. Considérations sur des Lépidoptéres envoyés du Guaté-
mala a M. de l’Aorza. Rennes.

108 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Butter, ArrHurR G. 1884. On a new species of the theclid genus Theritas from
Columbia. Ann. Mag. Nat. Hist. (Ser. 5) 14: 267.

Ciencu, Harry K. 1963. <A synopsis of the West Indian Lycaenidae with remarks
on their zoogeography. Jour. Res. Lepidoptera 2: 247-270.

Cramer, P. 1775. Uitlandsche Kapellen Waereld-Deelen. Vol. 1. Amsterdam.

Comstock, W. P. Anp E. I. Huntincron. 1958-1964. An annotated list of the
Lycaenidae (Lepidoptera, Rhopalocera) of the Western Hemisphere. Jour. N.Y.
Ent. Soc. 66: 103-118; 67: 59-95, 163-212; 70: 39-46, 100-118, 177-179;
71: 45-57, 72: 62-64, 120-130, 173-192.

Draupt, M. 1921. American Rhopalocera. In Seitz, A. Macrolepidoptera of the
World. Vol. 5. Stuttgart.

Druce, H. H. 1907. On Neotropical Lycaenidae with descriptions of new species.
Proc. Zool. Soc. Lond. (1907) pp. 566-631.

Fasricius, J. C. 1781. Species Insectorum. Vol. 2. Hamburg.

FreLper, C. anD R. 1867. Reise der Osterreicheschen Fregatte Novara um die Erde.
Vol. 2. Wien.

Gopman, F. D. anp O. Satvin. 1897-1901. Biologia Centrali-Americana. Insecta.
Lepidoptera-Rhopalocera. Vol. 2. London.

Hewirson, W. C. 1877. Illustrations of Diurmal Lepidoptera: Lycaenidae. Vols.
1, 2. London.

Husner, JAcos. 1819. Verzeichniss bekannter Schmetterlinge, Augsburg.

1827. Zutrage zur Sammlung exotischer Schmetterlinge. Vol. 5. Augs-
burg.

Kirpy, W. F. 1879. Catalogue of the collection of diurnal Lepidoptera formed
by the late William Chapman Hewitson. London.

Latuy, Percy I. 1930. Notes on South American Lycaenidae with descriptions of
new species. Trans. Ent. Soc. Lond. 78: 133-137.

Swarnson, WiLtiAM. 1832. Zool. Illus., 2nd Series, Insects. London.

Westwoop, J. O. 1852. Genera of Diurnal Lepidoptera. Vol. 2. London.

AN UNUSUAL VARIANT OF COLIAS PHILODICE (PIERIDAE)
IN NEW HAMPSHIRE

On October 5, 1970, I took a male Colias philodice philodice (Watreille), in
Lebanon, Grafton County, New Hampshire. This male is lacking all traces of the
smaller of the two ocelli on the underside of the hind wings with the exception of a
minute dot of color on the left. A closer examination of the ocelli showed that the
larger ocelli were incomplete where they joined vein Ms. An examination of the
uppersides of the hind wings showed that the corresponding orange colored spots

were incomplete, with the exception of a minute dot of color on the left side.
Due to the completeness of all other philodice markings, and also due to the
presence of part of the characteristic rings around the parts of the ocelli that are

present, there is little or no possibility of this specimen being mistaken for Colias
interior interior (Scudder. ) For reference purposes, this specimen has been deposited
in the collection of the Dartmouth College Museum (specimen # DCM-170-37-

19355 )

HARD I. Gray, Associate Curator of Biology, Dartmouth College Museum,
New Hampshire.

VoLUME 25, NuMBER 2 109

THE LIFE HISTORY OF SCHINIA CUPES DESERTICOLA
(NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute
Canada Department of Agriculture, Ottawa, Ontario

Schinia cupes Grote (1875, p. 113) is distributed from central Texas
(Georgetown, Irving) westward to the San Joaquin Valley of California
(Shafter, Coalinga), and northward in the Great Basin area to southern
Idaho (Twin Falls). On the deserts of southern California it is repre-
sented by a paler and less strongly maculate form named deserticola by
Barnes and McDunnough (1916, p.5). All of the Texas specimens exam-
ined as well as those from the San Joaquin Valley and from southern
Idaho are of the typical dark form.

The species is very common on both the Colorado and Mojave Deserts
of California, and all of the hundreds of specimens that have been exam-
ined from this area are of the pale vicariant deserticola. In the rather
limited series of genitalic slides examined, the valve of deserticola is some-
what narrower and the vesica somewhat shorter than in typical cupes.
The consistently paler colouring, the less intense maculation, and the
slight but possibly constant differences in male genitalic structure may
indicate that deserticola is actually a distinct though closely related spe-
cies. A knowledge of the habits and food plant of typical cupes will un-
doubtedly help to clarify the problem. For the present, it seems prefer-
able to retain the existing nomenclatorial status of deserticola as a
subspecies of cupes.

The Texas specimens of cupes were taken in April and May, the south-
ern Idaho specimens in mid June, and the San Joaquin Valley specimens
in April. Specimens of deserticola examined were taken during March
and April. In the spring of 1955, the flight at Thousand Palms in the Colo-
rado Desert of California was found to extend from 9 March to 10 April.
Three specimens taken at Indio, California on 28 October suggest that
pupal diapause in the species may be terminated by the occasional fall
rains that occur on the California deserts.

Behaviour

Schinia cupes deserticola feeds in the larval stage on the flowers and
seed capsules of Oenothera clavaeformis Torr. and Frém. (Figs. 2, 3).
Unlike many species of Schinia that are protectively coloured in their re-
semblance to the flowering heads of their food plants, the pale greyish-

110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

fawn colouring of adult deserticola bears no resemblance to the reddish-
pink flowering head of Oenothera clavaeformis. During a period in which
the heads of the Oenothera were being examined daily for eggs and young
larvae, no adults were found resting in the plants. The moth is evidently
exclusively nocturnal and the eggs are deposited at night, usually on the
still tightly closed buds (Fig. 2). A single wild-caught female deposited
a total of 93 eggs.

The newly hatched larva makes its way to the base of the bud and
bores through the calyx and corolla to gain entrance to the interior. Both
first- and second-stadium larvae feed preponderantly on the fleshy red
tissue of the inner surface of the receptacle. This habit is evidently re-
sponsible for the pinkish colouring that the larva assumes after a period
of feeding. During the third stadium, the larva attacks other sexual parts
of the plant. In the fourth stadium the larva usually quits the bud or
blossom and feeds from an exposed position on the flowering stem. In
the latter part of the fourth stadium and during the fifth, the larva feeds
almost exclusively on the younger seed capsules, and the buds are largely
abandoned as food. Unlike Schinia felicitata and Schinia florida which
bore through the wall of the seed capsule of other species of Oenothera
to eat the developing seeds, Schinia cupes actually consumes the whole
capsule. The more common red and mauve larvae are probably afforded
protection from predators by their general resemblance to the pink
flowering head of the food plant. All of the individually reared larvae of
deserticola matured in a uniform five stadia.

The larva tunnels into the soil to form a pupal cell and it is as a pupa
in the ground that the species spends the majority of the year.

Description of Stages

The following descriptions of immature stages are based on the progeny
of two females of deserticola taken at Thousand Palms, Riverside Co.,
Calif.; the description of the adult applies only to cupes deserticola.
Rearing techniques employed were those outlined by Hardwick (1958).
The duration of stadia listed are those for larvae reared at room tempera-

ture; the estimate of variability, following the mean for various values,
is the standard deviation.

Adult (Fig.

: |). Head and thorax grey irrorate with brown; abdomen pale fawn-
ye OW,

v. Forewing light fawn-grey. Transverse anterior line white with a dark inner
margin, consisting of three shallow excurved arcs. Basal space light fawn and con-
taining a variably expressed white or pale-grey basal line. Transverse posterior line
hite broadly excurved around cell, then essentially straight to inner margin; t.p.

| cakly scalloped between veins. Median space light grey irrorate with light
\ renzorm and orbicular spots large and prominent, and both with a

VoLUME 25, NuMBER 2 1B BD:

5

Figs. 1-5. Schinia cupes deserticola B. & McD. and its food plant. 1, Adult,
Thousand Palms, Calif.; 2, eggs on buds of Oenothera clavaeformis Torr. and Frém.;
3, Oenothera clavaeformis in its typical desert habitat; 4, dorsal aspect of ultimate-
stadium larvae; 5, ventral aspect of pupae.

dark central shade; orbicular always circular. A dark median shade or line passing
from costa through reniform and then paralleling t.p. line to inner margin. Sub-
terminal line an irregular pale shade, often with intervenal dark sagittate marks
proximal to it. Subterminal space fawn. Terminal space usually paler than sub-
terminal space. A series of intervenal dark-brown, marginal lines. Fringe checkered
fawn and brown. Hind wing dull cream with a broad, smoky-brown, outer-marginal

112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

4

S
jee)

Figs. 6-9. Schinia cupes desertico . & McD., ultimate-stadium larvae. 6, 7,

Left lateral; 8, 9, dorsal.

band. A pale patch usually evident in median area of outer-marginal band. A brown,
discal lunule usually prominently expressed. Veins often outlined by brown scales.
Fringe white variably marked with light brown, most strongly so at base. Underside
of both wings dull pallid cream marked with brown. Forewing with a prominent
brown reniform, a small dot-like orbicular, a variably expressed post-median line and
a paler, smoky-brown subterminal band. Fringe cream. Hind wing with a brown
discal lunule, a variably expressed, usually broken, post-median line and an evanes-
cent outer-marginal band. Fringe cream or white.

Expanse: 29.4 + 1.2 mm (100 specimens ).

Egg. Pale creamy-yellow when deposited and remaining so for the next 24 hours;
assuming a slight pink or orange tone on anterior half during second day. Pink tone
more pronounced and sharply defined on third and fourth days after deposition.
Whole egg turning dark grey with head and prothoracic shield becoming visible
through chorion a few hours before hatching.

Dimensions of egg: length, 0.52 + 0.07 mm; diameter, 0.64 + 0.03 mm (5 eggs).

Duration of egg stage: 4.4 + 0.6 days (97 eggs).

Virst-Stadium Larva. Head very dark brown or black. Prothoracic and suranal
shields dark smoky-brown. Trunk creamy-white or greyish-white and often stained
with pink. Thoracic legs and proleg shields smoky-brown.

Head width: 0.31 + 0.01 mm (25 larvae).

Duration of stadium: 3.6 + 1.0 days (21 larvae). .

Second-Stadium Larva. Head capsule light orange-brown variably suffused and
mottled with medium brown. Prothoracic shield pale fawn marked with brown along
margins. Suranal shield pale fawn lightly marked with greyish-brown. Trunk green-
ish-grey or yellowish-grey and often stained with pink; dorsum of trunk with a
median and a pair of subdorsal light reddish-brown lines; a pale-cream or greyish-
piracular band. Thoracic legs and proleg shields dark smoky-brown.

0.53 + 0.03 mm (25 larvae).

VoLUME 25, NuMBER 2 ALS}

Figs. 10, 11. Schinia cupes deserticola B. & McD., apical abdominal segments
of pupa. 10, Ventral; 11, right lateral.

Duration of stadium: 2 days (21 larvae).

Third-Stadium Larva. Head light fawn, weakly mottled with light chocolate-
brown. Prothoracic shield pale fawn tinged with pink or green. Suranal shield pink
or green depending on body colour. Trunk varying from cherry-red through pink and
various shades of brown to leaf-green. Mid-dorsal band usually the darkest area of
trunk. Subdorsal area paler than mid-dorsal band, and with poorly defined marginal
lines of white or cream. Supraspiracular area only slightly paler than mid-dorsal
band. Spiracular band light yellow in green specimens, cream or white suffused
with pink in red and brown specimens. Suprapodal area concolorous with subdorsal
area. Mid-ventral area grey, greenish-grey or pinkish-grey. Thoracic legs cream or
very pale fawn.

Head width: 0.92 + 0.04 mm (25 larvae).

Duration of stadium: 2.1 + 0.5 days (21 larvae).

Fourth-Stadium Larva. Head cream or pale fawn variably mottled with darker
fawn. Prothoracic shield poorly distinguished and suranal shield undistinguished
from remainder of trunk. Trunk cherry-red, purplish-brown, chocolate-brown or
green. Mid-dorsal band usually somewhat darker than subdorsal area but often poorly
distinguished from it; in some cases mid-dorsal band margined by evanescent pale
lines. Subdorsal area separated from supraspiracular area by a pale greyish-cream
line. Supraspiracular area somewhat darker than mid-dorsal band, lightly flecked
with cream or grey. Spiracular band yellowish-cream. Suprapodal area concolorous
with or somewhat paler than subdorsal area. Mid-ventral area dull grey suffused
with colour of body. Spiracles with medium brown rims. Thoracic legs pale fawn.

Head width: 1.63 + 0.06 mm (25 larvae).

Duration of stadium: 4.4 + 1.0 days (21 larvae).

Fifth-Stadium Larva (Figs. 4, 6-9). Head light orange-brown with cherry-red
reticulations and markings. Prothoracic shield fawn, usually heavily suffused with
red or mauve, usually poorly distinguished from trunk. Suranal shield red or mauve,
essentially indistinguishable from remainder of trunk. Mid-dorsal band varying from
medium chocolate-brown to light red, often paler near posterior end of each segment
so that it has a broken appearance. Subdorsal area varying from mauve to red,
sometimes well-defined from mid-dorsal band, sometimes almost indistinguishably
fused with it. Supraspiracular area darker than subdorsal area or concolorous with
it; occasionally suffused with red or spotted with brown and often with white arcuate
marks. Spiracular band light yellow or cream. Suprapodal area cherry-red or mauve,

114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

paler than dorsal region, also with white arcuate marks. Mid-ventral area grey, some-
times suffused with pink. Spiracles with dark-brown rims. Thoracic legs pale fawn
or cream.

Head width: 2.60 + 0.10 mm (16 larvae).

Duration of feeding phase of fifth stadium: 5.3 + 1.7 days (21 larvae).

Duration of prepupal phase of fifth stadium: 3.3 + 1.0 days (12 larvae).

Pupa (Figs. 5, 10, 11). Orange-brown. Spiracles on a level with general surface
of cuticle or borne in very shallow depressions. Rims of spiracles only weakly project-
ing. Anterior areas of abdominal segments 5, 6 and 7 conspicuously but rather finely
pitted. Apex of proboscis terminating between apexes of forewings. Cremaster con-
sisting of two elongate, slender, slightly curved spines borne at the apex of a rounded
prolongation of tenth abdominal segment.

Length from anterior end to posterior margin of fourth abdominal segment: 11.1 +
0.6 mm (17 pupae).

Acknowledgments

I am grateful to Mr. John E. H. Martin of this Institute for the fine
photographs accompanying this paper and for his assistance in the field.
I appreciate the help of my associate, Mr. E. W. Rockburne, who mea-
sured the immature stages and drew the cremaster area of the pupa.

Literature Cited

Barnes, W., AnD J. H. McDuNNoucH. 1916. Contributions to the Natural History
of the Lepidoptera of North America. 3(1): 5. Decatur, IIl.

Grote, A. R. 1875. Descriptions of North American moths. Trans. Amer. Ent.
Soc. 5: 113-118.

Harpwick, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae) with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

THE PAINTED LADY BUTTERFLY, VANESSA KERSHAWI
(NYMPHALIDAE), OF AUSTRALIA AND NEW ZEALAND

Williams (1970, J. Lepid. Soc. 24: 157-175) refers to the Painted Lady Butter-
fly of Australia, New Zealand and some Pacific Islands as a form kershawi of Vanessa
cardui (Linn.). He says that this form has slight differences in structure and wing
markings, and has sometimes been considered as a distinct species.

[ would like to point out that van Son (1966, J. ent. Soc. Od 5: 66) has shown
Vanessa kershawi (McCoy) to be a distinct species on the basis of genitalia. There
are also constant differences in wing markings which support the separation of
kershawi from cardui.

tecent migrations of V. kershawi in Australia have been recorded by Smithers
/

and Peters (1966, J. ent. Soc. Od 5: 67-69) and Smithers (1969, Aust. Zool. 15

) 12R 1
| } £OO— Jl 4).

}. VY. Perens, The Australian Museum, Sydney, N.S.W., Australia.

VOLUME 25, NUMBER 2 BIS

BIOLOGICAL AND SYSTEMATIC CONSIDERATIONS ON THE
“EMARGINANA GROUP” OF THE GENUS EPINOTIA
(TORTRICIDAE)!

PauL A. OPLER?
University of California, Berkeley

In his revision of the Eucosminae, Heinrich (1923) considered the
three Nearctic members of the genus Epinotia Hiibner with an emar-
ginate termen of the forewing to constitute a distinctive species group.
In his treatment he gave only a few features required to separate the
species, presented photographs of genitalia preparations which do not
illustrate diagnostic features, and gave no biological information other
than host records.

In this paper I will illustrate the male genitalia of these three species,
and give additional distributional and biological information for the group
based on material in the California Insect Survey, University of California,
Berkeley, California. Discussions of the biological features of E. emar-
ginana and E. crenana are based on notes taken by J. A. Powell and the
author. In addition, a lectotype is selected for Epinotia emarginana
(Walsingham).

Epinotia emarginana (Walsingham )
Proteopteryx emarginana Walsingham, 1879, Illus., Lepid. Heter. Brit. Mus., 4:68.
Epinotia emarginana; Heinrich, 1923, U.S. Nat] Mus. Bull. 123:218-219; MacKay,

1959, Can. Ent. Supp. 10:113; Powell, 1962, Pan-Pac. Entomol. 38:134; Powell,

1964, Univ. Calif. Publ. Entomol. 32:33, 51.

The late N. S. Obraztsov examined the type series of this species in the
British Museum (Natural History), and selected a lectotype, but he did
not publish this designation. In order to fix the specific identity of this
species I designate the specimen indicated by Obraztsov to serve as the
lectotype of this species. This individual is a male syntype and bears the
following data: “above Potter's Valley, Mendocino Co., California 13.
VI 1871 Wlsm slide 11604.” A photograph of the genitalia of the lectotype
is shown as Figure 7.

This “species” presents a number of perplexing problems. All of these
relate to the possibility that two sibling species are included under the
name emarginana. The information which indicates this possibility is
biological and will be presented after a discussion of the morphological
features of emarginana.

1 Funds supporting field and laboratory work during this study, conducted under the direction

of J. A. Powell, were provided by National Science Foundation grants GB 4014 and GB 6813X.
2 Current address: Organization for Tropical Studies, Universidad de Costa Rica, Costa Rica.

116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-6. Examples of Epinotia emarginana polymorphs. 1, 2, Males, Straw-
berry Canyon, Alameda Co., Calif., 16 April 1960, r. f. Quercus agrifolia (J. Powell);
3, male, Applegate vic., Placer Co., Calif., 24 April 1961, r. f. Quercus kelloggii (J.
Powell); 4, female, Orinda, 1 mile north, Contra Costa Co., Calif., 19 April 1967,
r. f. Quercus agrifolia (P. Opler); 5, female, Fairfax, 2 miles south, Marin Co., Calif.,
30 April 1967, r. f. Quercus agrifolia (P. Opler); 6, female, Applegate vic., Placer
Co., Calif., 24 April 1961, r. f. Quercus wislizenii (J. Powell).

Morphologically this species differs from the other two members of
the complex in maculation features, scaling of the labial palpi, and char-
acteristics of the male genitalia.

The coloration of the legs, hind wings and abdomen is relatively in-
variable. The legs have the femur tan scaled. The pro- and mesothoracic
tibiae are brown with a few tan scales intermixed, while the mesothoracic

| are pale tan. The tarsomeres are brown basally and edged with

Pr ylive

VoLuME 25, NuMBER 2 REZ

pale tan distally. The hind wing is brown. The abdomen is brown dor-
sally and pale gray-tan ventrally.

While the above features are relatively constant, the coloration of the
head, dorsal surface of the thorax, and particularly that of the forewings
is extensively polymorphic. The color and pattern of these latter features
appear to be determined by a number of independently operating genes
or sets of genes. Thus there appears to be no definable set of polymorphs.
In fact, it is almost impossible to find two individuals with nearly iden-
tical wing color and pattem. The ground color varies from tan, gray,
various shades of brown, chestnut to black; the wing pattern may be
obsolescent, with a large patch on the inner margin, several patches in
the wing median, or a general mosaic of various colors. Several frequent
morphs are shown in Figures 1-6. Throughout this melange of variation
the pattern of the costal margin, apex, and outer margin tends to remain
constant. The costal margin usually has seven white, cream, or silvery
outwardly angled marks, each of which is divided by a small patch of
dark scales. On the posterior half of the forewing just in from the outer
margin are two parallel patches of scales which appear silvery in reflected
light. The fringe on the outer margin is of white-tipped brown scales.

Because of the pattern of scaling, the labial palpi of E. emarginana
are distinctively shaped when viewed laterally. The dorsal margin is con-
vexly curved, the outer margin truncated, and the ventral margin straight
(Fig. 13).

The male genitalia of E. emarginana are characterized by the absence
of a gnathos, by the recurved arms of the socii, which are dentate dor-
sally, by the blunt uncus, by the neck incurvation (sensu Heinrich, 1923)
at the ventral margin which extends half the width of the valva and by
the rounded valval tips ( Fig. 16).

The female genitalia are illustrated in Figure 19; the female genitalia
of the other two species differ little from that of E. emarginana and are
not illustrated.

Biological information suggests that two sibling species may be in-
volved in populations presently referred to emarginana, although inten-
sive morphological comparisons have not supported this thesis. Of the
two putative entities one utilizes species of Quercus (Fagaceae) as hosts,
while the other feeds on Arctostaphylos and Arbutus (Ericaceae).

The life cycles are similar for both host types, but apparently are tem-
porally out of phase. Larvae are found on Quercus primarily during April
but have been found as early as 16 March and as late as 10 May. During
this time the larvae show a preference for the flowering parts, but will

118 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

also feed on young foliage. Larvae associated with Arctostaphylos were
collected on young foliage between 19 May and 6 June. The larvae on
Arbutus represent a single lot collected in flowers of Arbutus menziesii
Pursh on 14 April.

Upon reaching maturity the larvae drop from the hosts and pupate.
The cocoons are covered with detritus and soil particles. Eclosion occurs
about three weeks later; moths originating from oak emerging during
April and May, those from manzanita, during June.

The newly emerged adults are not functionally reproductive and
possess an inordinate amount of fat body material. Adults collected in the
fall still display this pre-reproductive condition. Beginning in January,
six to eight months after eclosion, and continuing into April mated fe-
males may be found in the field. At this time males are rare, although
the collection of a pair in copula on 6 March supports the thesis that mat-
ing does not occur until after winter has passed. Eggs are laid on twigs
or leaves of the host. Females collected on 8 January and 6 February
laid eggs while caged in the laboratory.

Presumably if two species are involved, differential timing of reproduc-
tive maturation, and mating could serve as an isolating mechanism. How-
ever, if mating and oviposition of a single species is occurring in a con-
tinuum, the early developing females presumably find Quercus to be
preferrable oviposition substratum while late developing females find
Arctostaphylos to be the suitable oviposition substratum.

The following constitute rearing records for this species based on ma-
terial in the California Insect Survey. Data for adults reared from Faga-
ceae and Ericaeae are listed separately.

Records from Fagaceae. Alameda Co.: Berkeley, IV-14-58 r. f. Q. agrifolia,
19 emgd. V-5-58 (J. Lannon); Strawberry Cyn., IV-16-60, IV-6-67, IV-11-68 r. f.
Q. agrifolia JAP 60D3, 67D15, 68D126, 496 ¢ 449 2 emgd. V-8/15-60, V-10-67,
V-5/18-68 (J. Powell, P. Opler). Contra Costa Co.: Antioch, 2 mi. E., IV-8-67 r. f.
QO. agrifolia JAP 67D61, 16 emgd. V-4-67 (P. Opler); Orinda, 1 mi. N., IV-19-67,
[V-9-68 r. f. Q. agrifolia, Q. kelloggii, JAP 67D75, D 81, 68D87, 62 6 42 2 emed.
V-15/16-67, V-5/14-68 (P. Opler); Russell Farm, 4 mi. NE Orinda, IV-7-67, IV-8-
67 r. f. QO. agrifolia JAP 67D15, 68D105, 54 ¢ 49 9 emgd. V-12-67, V-3/12-68 (P.
Opler); Walnut Creek, V-10-67 r. f. Q. lobata JAP 67E21, 19 emgd. V-30-67 (J.
Powell). Fresno Co.: Kings Canyon Nat’] Prk., IV-16-60 r. f. oak in chapparal, 12
emgd., V-19-60 (W. E. Ferguson). Glenn Co.: Elk Crk., 10 mi. NW, IV-21-68
r. f. O. dumosa, Q. wislizenii var. frutescens JAP 68D167-168, 24 ¢ 229 emegd.
V-16/20-68 (P. Opler). Marin Co.: Fairfax, 2 mi. S, IV-29-67 r. f. QO. agrifolia
JAP 67D134, 16 19 emgd. V-29/VI-2-67 (P. Opler); Inverness, 2 mi. SE, IV-13-
63 r. f. O. agrifolia JAP 68D130, 14 19 emgd. V-16/18-68 (P. Opler). Placer Co.:
\pplegate, vic., IV-24-61 r. f. Q. kelloggii, QO. wislizenii JAP 61D9, 26 4 39 2 emed.
V-18/24 61 (J. Powell); Rocklin, IV-18-68 r. f. QO. wislizenii JAP 68D142, 34 4

2 em: d /-12/18-68 (P. Opler). San Francisco Co.: Golden Gate Park, IV-
75-68 QO. agrifolia JAP 66D15, 68D50, 26 6 emgd. V-15-66, V-18-

VoLUME 25, NuMBER 2 119

Fig. 7. Male genitalia from lectotype of Proteopteryx emarginana Wlsm.

Figs. 8-11. Polymorphs of Epinotia crenana. 8, Female, Berkeley Hills, 1400’,
Contra Costa Co., Calif., 15 April 1968, r. f. Salix sp. (P. Opler); 9-11, male and
two females, Pacific Grove, Monterey Co., Calif., 15 April 1962, r. f. ornamental
Salix (J. A. Chemsak).

Fig. 12. Epinotia cercocarpana, female, Crook Creek lab., 10, 150’, White
Mountains, Mono Co., Calif., 4 July 1961, r. f. Cercocarpus ledifolius (J. Powell).

68 (P. Opler). Santa Barbara Co.: Santa Cruz Isd., Prisoner's Harbor, III-16-69
r. f. Q. agrifolia JAP 68C52, 106 6 922 emgd. IV-10/27-69 (P. Opler & J.
Powell). Shasta Co.: O’Brien, Shasta Lk. Rec. Area, JAP 68D159, D161, 64 ¢
522 emgd. V-16/23-68 (P. Opler); Project City, IV-20-68 r. f. Q. douglasii, Q.
lobata JAP 68D162-3, 26 ¢ emgd. V-16-68 (P. Opler). Tehama Co.: Redding,
10 mi. E, IV-20-68 r. f. OQ. wislizenii JAP 68D156, 1¢ emgd. V-18-68 (P. Opler).

Records from Ericaceae. Contra Costa Co.: Berkeley Hills, IV-14-66 r. f.
Arbutus menziesii JAP 66D18 emgd. V-9-66 (J. Powell & J. Wolf). Marin Co.:
Alpine Lake, VI-4-56, VI-6-57 r. f. Arctostaphylos sp. JAP 57F8 emgd. VI-27-56,

120 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

x
ZY a es
AISNE ee 2 eS Fe, ~\
WINK =i a3 SLeS RS

N\A > aS 2S ’ Ras a EN ae

SS SSS SN See

= 9 ~> > :

SS53 SS pecs

Figs. 13-15. Schematic view of heads of Epinotia spp. showing shape of labial
palpi. 13, E. emarginana; 14, E. crenana; 15, E. cercocarpana.

VII-1-57 (J. Powell). Mendocino Co.: Booneville, vic., VI-1-57 r. f. Arctostaphylos
JAP 57F1 emegd. VI-21/VII-1-57 (J. Powell); Leggett, V-19-66 r. f. Arctostaphylos
JAP 66E17, emged. VI-14/23-66 (J. Powell & J. Wolf).

Epinotia crenana (Hubner)

Tortrix crenana Hubner, 1827, Samm. Eur. Schmett. Tort. Fig. 242.

Epiblema crenana; Staudinger and Rebel, 1901, Cat. Lepid. 2, #2133.

Eucosma crenana; Dyar, 1904, Proc. Ent. Soc. Wash. 6:117.

Proteopteryx columbia Kearfott, 1904, Can. Ent. 36:112.

Proteopteryx columbia mediostriana Kearfott, 1904, Can. Ent. 36:114.

Epinotia crenana; Heinrich, 1923, U.S. Natl Mus. Bull. 123:219; MacKay, 1959,

Can. Ent. Supp. 10:110; Powell, 1962, Pan-Pac. Entomol. 38:134.

This Holarctic species possesses a range of polymorphism less extensive
than that displayed by E. emarginana. E. crenana lacks the distinct
costal marks and reflective patches of E. emarginana. Dull browns and
tans are the colors most often shown by the various morphs, while white
and reddish are generally lacking. One macular feature which is fre-
quently present in crenana, but lacking in the other two species, is a thin
black streak on the costal half of the forewing two-thirds of the distance
from the base. Some typical morphs of crenana are shown in Figures 8-
i

The labial palpi of E. crenana when viewed laterally have the dorsal
and ventral margins flaring and truncated distally (Fig. 14).

Genitalically this species is quite distinct (Fig. 17). In addition to the
invasion of the cucullus by the neck incurvation mentioned by Heinrich
(1925, p. 198), the uncus is notched with two acuminate projections dis-
tally, the socii are triangulate and the gnathos is strongly reduced.
| he larvae of crenana feed exclusively on Salix species (Salicaceae).

lary ae are fom

id in shelters on young foliage or as inquilines in the

VoLUME 25, NUMBER 2 oA

Figs. 16-18. Male genitalia of Epinotia spp. 16, E. emarginana. (PAO prep.
115); 17, E. crenana. (JAP prep. 49); 18, E. cercocarpana. (JAP prep. 2463).

122 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 19. Female genitalia of Epinotia emarginana. (PAO prep. 65).

leaf and petiole galls of tenthredinids. Larval collection dates, which

range from 8 March to 17 September, indicate that crenana is multiple-
brooded.

Distribution. Holarctic; British Columbia and Manitoba constituted the known
distribution in North America at the time of Heinrich’s revision (1923). The Cali-
fornia Insect Survey has reared material from several localities in California and
Nevada as follows: California—Alameda Co.: Berkeley, Univ. Calif. Campus,
IJ-11-58 r. f. Salix JAP 58C7 emgd. IV-8-58 (J. Powell). Contra Costa Co.:
Berkeley Hills, 1400’, NE Oakland, IV-15-68 r. f. Salix sp. JAP 68D137 emgd.
V-18/23-68 (P. Opler); El Sobrante, III-8, IV-26-58 r. f. Salix JAP 58C2, D13,
emgd. V-8/26, V-26/VI-9-58 (J. Powell). Monterey Co.: Pacific Grove, IV-15-62
r. f. ornamental Salix JAP 62D9 emgd. by V-22-62 (J. A. Chemsak). Los Angeles Co.:
Pacific Palisades, V-56 r. f. Salix leaf galls (W. Hovanitz). Santa Clara Co.: San
Jose, IV-16-61 r. f£. Salix lasiolepis leaf gall emgd. VI-25-61 (L. R. O’Brien). Nevada.
—Washoe Co.: 2 mi. SW Reno Hot Spg., 6100’, IX-17-65 r. f. Euura petiole gall
on Salix lasiandra emgd. X-28-65 (E. L. Smith).

Epinotia cercocarpana (Dyar)

Eucosma cercocarpana Dyar, 1903. Proc. Ent. Soc. Wash. 5:297.
Epinotia cercocarpana; Heinrich. 1923. U.S. Natl Mus. Bull. 123:219.

I’. cercocarpana is characterized by its nonpolymorphic, mottled with
black gray appearance (Fig. 12). The vestiture of the head, thorax and
ng composed of white, black, and white-tipped black scales.

VoLUME 25, NuMBER 2 8}

The forewing maculation is somewhat variable, but cannot be termed
polymorphic. There is usually a small white triangular patch on the
costal margin at the apex, a triangular black or dark gray patch on the
inner margin near its junction with the outer margin, and an outwardly
directed black bar beginning on the costal margin halfway from the base
and extending nearly to the triangular black patch. In five of seven
specimens the scaling of the thoracic tergites is black on the posterior
half and gray on the anterior half.

In lateral view the labial palpi appear to be convexly curved dorsally
and slightly flaring ventrally due to the pattern of scaling (Fig. 15).

In addition to triangular socii mentioned by Heinrich (1923, p. 198),
there are several other distinctive features in the male genitalia of the
moth (Fig. 18). The gnathos arms are separate, narrow, and strongly
recurved, the neck incurvation of the valva is the shallowest of any of the
species in this group and the valvae are truncate distally.

Distribution. The species was previously known only from the type of locality,
Platt Cyn., Douglas Co., Colorado. New distributional data are as follows: Ari-
zona.—Mojave Co.: Hualpai Mtn. Park, VI-2-68, at B/W light (P. Opler & J.
Powell). California.—Inyo Co.: Crooked Crk. Lab., 10,150’, White Mtns. VII-4-61
r. f£. Cercocarpus ledifolius JAP 61G1.1 emgd. VIII-31-61 (J. Powell). Tuolumne Co.:
Strawberry, VIII-20-60 (D. Q. Cavagnaro). Colorado.—El Paso Co.: Rock Crk.

Cyn., Colorado Springs VII-16-59, VII-15-60 (Margot May). Jefferson Co.: Golden,
X-25-67 (no collector, Bryant Mather Collection ).

Acknowledgments

I wish to thank F. H. Rindge, American Museum of Natural History,
New York, for loaning the photographs of type material for Epinotia
emarginana made by the late N. S. Obraztsov.

J. A. Powell and G. R. Rotramel, University of California, Berkeley,
provided early insight into the present complex and reviewed the manu-
script of this paper.

My wife, Sandra, gave welcome secretarial assistance.

Literature Cited

HernricH, Cart. 1923. Revision of North American Moths of the subfamily
Eucosminae of the family Olethreutidae. U.S. Nat’ Mus. Bull. 123.

124 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

A MIGRATION OF KRICOGONIA CASTALIA (PIERIDAE)
IN NORTHERN MEXICO?

GrEoRGE W. BYERS

Dept. of Entomology, University of Kansas, Lawrence

Kricogonia castalia Fabricius ranges from northern South America
(Venezuela) through Central America and Mexico into Texas and
through the Caribbean region (Haiti, Puerto Rico, Jamaica) occasionally
to southern Florida. It has been reported (often under the synonymous
name, K. lyside Latreille) as common in Texas, and stray individuals
have been recorded as far north as Colorado, Nebraska and Illinois ( Ehr-
lich and Ehrlich, 1961). According to Godman and Salvin (1887-1901,
p. 150), K. castalia (as K. lyside) “does not appear to be a very common
insect” in Mexico and Central America.

A migratory flight of this species, observed in central Tamaulipas on
11 July 1961, suggests it sometimes becomes at least locally abundant in
northern Mexico. The butterflies were seen moving generally from
northwest to southeast, toward the Gulf of Mexico. These observations
were made about 80 miles inland from the Gulf, along Highway 101,
commencing about 14 miles northeast of Ciudad Victoria near Giiémes.
From there, the width of the stream of Kricogonia was measured as 53.5
miles on to the northeast. I have no estimate of the length of the emigrat-
ing stream. Since the terrain along this part of the highway is rather flat
with only light and scattered vegetation, the butterflies, especially when
moving in dense groups, could be seen several hundred yards to either
side of the road. Their bright yellow on pale yellow color also made them
easy to see from afar.

_ Density of the migration varied within the broad zone described, yet
tor the entire 53.5 miles some butterflies were always in sight, moving
generally toward the southeast. The widest of the dense streams within
the migration was 9.5 miles across. Since the butterflies flew generally
within six feet of the ground, and mostly about two or three feet above
the surface, large numbers of them were struck and killed by our vehicles
as we drove across the emigrating stream at about 50 miles per hour. It
is not easy to estimate the density of insects in such an emigration, but
my counts made at the time suggest about 1100 butterflies per minute
were crossing an average mile of front along the highway (or perhaps
some 55,000 per minute were passing a line at right angles to the direc-

( nt ibutio 1 AF A . .
EO AU 1462 from the Department of Entomology, University of Kansas, Lawrence.

_R, Paylor for reading the original manuscript of this note and making
arly concerning dissection of females.

VOLUME 25, NUMBER 2 125

tion of the stream). A sample of 18 specimens was easily taken in a
couple of minutes.

At the time of these observations, the weather was hot and the sky was
partly cloudy, the gathering clouds producing a light sprinkle of rain ap-
proximately 65 miles northeast of Ciudad Victoria, near mid-afternoon,
just before we drove out of the migration. There was a light to moderate
breeze from east-northeast, so that the butterflies, on a northwest to south-
east course, were moving slightly into the wind.

In the sample collected, there was a 3.5 to 1 ratio of females to males
(142,44). Three females with large abdomens were dissected, and it
was found that none contained any eggs whatsoever (yet large amounts
of what appeared to be fat) and that each contained the hardened re-
mains of one large spermatophore. In the males, there was noticeable
variation in the extent of the black spot near the costal margin of the
hind wing. Adults of K. castalia have been seen feeding on blossoms of
mesquite, and Klots (1951) reports lignum vitae (Guaiacum officinale )
as larval food for this species in Puerto Rico. Unless this emigration had
come a long way, it seems the food plant must have been some common
member of the semi-desert flora.

Migrations of pierid butterflies are frequently observed and reported
in the entomological literature; however, as far as I can determine, this
is the first record of a migration of a species of Kricogonia (there are
only two species in the genus). Recent books on insect migration ( Wil-
liams, 1958; Johnson, 1969) do not mention the genus.

Literature Cited

Exruicu, P. R. anp A. H. Exriuicn. 1961. How to know the butterflies. W. C.
Brown Co., Dubuque, Iowa.

GopMaNn, F. D. anp O. Satvin. 1887-1901. Biologia Centrali-Americana. Lepidop-
tera-Rhopalocera, vol. 2.

Jounson, C. G. 1969. Migration and dispersal of insects by flight. Methuen &
Cos td. London.

Kuots, A. B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Bos-
ton.

WituiaMs, C. B. 1958. Insect migration. Macmillan Co., New York.

126 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

OBSERVATIONS ON PAPILIO ARISTODEMUS PONCEANUS
(PAPILIONIDAE)

FRANK RUTKOWSKI
153 Centre St., New York, N.Y.

Papilio aristodemus ponceanus Schaus is a swallowtail which few
Lepidopterists have seen alive. As the North American population of a
polytypic Antillean species, it is limited to tropical hardwood hammocks
in southern Florida. Before the extensive land clearing which accom-
panied settlement, its range may have extended from a southwestern
limit near Marathon in the Florida Keys, into peninsular Florida as far
north of Miami as soil character and climatic fluctuations would permit
these hammocks to flourish. Today its range is greatly reduced.

The life history was described by Grimshawe (1940) who also illus-
trated some of the early stages. The accompanying figures will supple-
ment her photographs.

During late spring of 1970, I made observations which can be added
to those of Grimshawe. I spent the first week of June in the habitat of P.
a. ponceanus at times of day varying from dawn to just before sunset.
During this time I watched about 35 ponceanus. Nearly half of these
were caught in order to determine sex and condition, but almost all were
set free. Except for a female released and then recaptured an hour later
and a mile away, it is not possible to say how many individuals these
sightings might represent.

Circumstances did not permit the rigorous survey with marked speci-
mens and large sampling which would have made these notes more ob-
jective. Nevertheless I offer this tentative account because ponceanus
is on the verge of extinction. Collectors hunt it down while developers
are destroying its habitat (Kimball 1965, Klots 1951). As I hope to
show, over-protective conservation practices can endanger it further.

Published dates for ponceanus (Henderson 1945a, 1945b, 1946) extend
mostly from April to June. Although the finding of two larvae in their
third instar on June 5 (discussed below) indicates that parents had been
on the wing in mid-May, the main 1970 flight seemed to have started
with the heavy rains which culminated in Hurricane Alma near the end
of May. Local newspapers claimed that this had been the wettest May
in years. And when I arrived on the afternoon of June 2, residents told
me about the unusually heavy and continuous rains which had stopped
ut the night before. The weather stayed mostly fair until the 5th, when

vilw |

‘ly late in the afternoon and intermittently through the

VOLUME 25, NUMBER 2 DAT

night. Between these two rains ponceanus was locally in fresh condi-
tion and not rare, whereas the usually common butterflies were scarce
and worn. After the rain on the 5th, the few ponceanus I could find were
all shabby, and ordinarily abundant species such as Nathalis iole Boisdu-
val, Junonia coenia (Hiibner) and Danaus gilippus berenice (Cramer )
first appeared as freshly-emerged singletons.

Adults. Ponceanus usually flew in brushy hammocks. The earliest
any were seen was just before 9 AM (Eastern Daylight Time) on two
bright days, in each instance a female visiting the blossoms of Cheese
Shrub (Morinda Roioc L.) at the eastern edges of hammocks. The latest
observation was at 5 PM, a freshly-emerged female hovering low over
Cheese Shrub blossoms at the sunlit west edge of open hammock. On
June 2, 1970, sunrise in the Miami area was at 6:29 AM (E.D.T.) and
sunset was at 8:08 PM.

Before the hottest part of the day, which was from 1 to 2 PM, both
sexes were within the hammocks, fluttering in diffused light about a foot
above the ground at blossoms of Guava (Psidium guajava Raddi). Guava
was abundant within the hammocks whereas Cheese Shrub grew com-
monly at the edges of hammocks or out in the open. Those ponceanus
seen at flowers within hammocks were invariably visiting Guava, while
at the edges of hammocks or out in the open they were visiting only
Cheese Shrub when they were on flowers at all.

On bright days the butterflies flew higher during the hot afternoon.
Males patrolled the tree tops at a height of 10 feet or more, sometimes
descending into open spaces to investigate any other ponceanus. How-
ever, no combats were observed nor was it possible to see whether in-
dividual males kept to flyways above the forest canopy. During the
heat of the day I rarely saw a male descend from the canopy to feed at
flowers.

At 10 oclock on a hazy morning after a drizzle, I found a male of
Papilio cresphontes cresphontes Cramer and a male ponceanus along an
open trail. After a short encounter the ponceanus veered into the woods
while the cresphontes settled on a shrub to bask. Three other cresphontes
were found in open places near midday; all were males visiting various
flowers in bright sunlight.

Regardless of how fresh they were otherwise, many ponceanus males
had one or both hind wings torn more or less cleanly at a 90° angle to the
anal margin, often with the tails intact. No females were found damaged
in this way. Such mutilation may be evidence of bird attack, although
no specimen bore a v-shaped beak mark. Although none of the four male
cresphontes had torn wings, one was missing a tail which apparently had

128 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

been pulled out from its root. The tails of cresphontes are spatulate,
marked above and below with a conspicuous yellow spot; ponceanus tails
are slender and dark. It is possible that the conspicuous cresphontes
tails might offer marginal survival advantage by distracting the attention
of birds from the body of the insect or from graspable parts of the wings.
Male ponceanus seemed to fly just as strongly as male cresphontes when
out in the open. I could not distinguish the two species by flight alone.

A ponceanus male and a cresphontes male of about the same size were
roughly compared for wing strength by measuring the diameter of each
wing vein with a micrometer at a point 4” from the outer margin. The
averages were then taken. The average reading for the ponceanus male
was .0165” while the cresphontes average was only .0150”. A female of
each species similarly compared showed less discrepancy. They both
averaged .0185”. Though the ponceanus male might seem to be the more
robust insect, other things being equal, it is worth mention that the
pointed shape of cresphontes wings increases the rigidity of wing surface
relative to vein diameter. From the standpoint of total mass that must be
kept in motion to maintain a given speed, to say nothing of maneuver-
ability, the pointed cresphontes wings seem to be more efficient than the
blunt ponceanus wings.

In an attempt to see how important wing pattern or color might be in
species recognition, I looked at a pair of cresphontes and a pair of ponce-
anus under long-wave ultraviolet light. The bulb used was a General Elec-
tric 8 watt BLB, which I hoped would approximate the wavelength by
which these insects perceive pattern or color. Under ultraviolet illumina-
tion the relation of light spots to dark background on the upperside of
both species was similar to the relation visible in daylight. The under-
sides of the cresphontes pair also looked much as they do in daylight.
However, the undersides of the ponceanus pair differed strikingly from
cresphontes. The wide orange-brown band on the hind wings which is
so Characteristic of ponceanus (but which tends to vanish in Antillean
aristodemus populations) was invisible under ultraviolet light. Fur-
thermore, the yellow underside ground color was nearly obliterated.
Ixxcept for the blue lunules on the hind wings, the entire underside of
the ponceanus pair appeared a deep violet brown. For this reason I
suspect that cresphontes and ponceanus can distinguish each other in
Hight better than a Lepidopterist can.

Living males of ponceanus smell strongly of sweet lemon with a sug-
gestion of vanilla, while a freshly-emerged female had no perceptible

ent. A living male cresphontes caught in the same place at the same

VoLUME 25, NuMBER 2 129

time, had a faint odor which is best described as acrid. These scents fade
soon after death.

At 10:15 AM in a dimly-lit trail through brushy hammock I saw a
female and two male ponceanus visiting flowers at opposite ends of a
Guava tangle. The fresher of the two males eventually approached this
slightly worn female while she was still feeding. He hovered over her.
She then settled on the ground with wings flattened and vibrating, raising
her abdomen. The male fluttered on the ground behind and then rose
over her before flying away. In contrast, during part of a cresphontes
courtship witnessed at 3 PM on a bright day in May 1966, the pair flew
straight up until they were hovering about 30 feet above nearly bare
ground. Their nuptial flight was interrupted by a bird which darted up
out of nearby mangroves to attack one of the butterflies from a “five
oclock” position. The cresphontes evaded this attack by an easy dip
sideways while its would-be mate fled.

Female ponceanus behaved differently from males. The few times
they were seen along the edges of hammocks their sex could seldom be
determined unless they were captured. But when flying within ham-
mocks in search of Torchwood, Amyris elemifera L., the larval foodplant,
they fluttered slowly about 6 feet from the ground, with the axis of the
body at about 45° from horizontal.

Oviposition. Several ponceanus females at the outer edges of ham-
mocks were seen to investigate shrubs of Wild Lime, Zanthoxylum Fagara
(L.), a common larval foodplant of cresphontes in southern Florida.
These females all flew lower than usual, at about 4 feet. One female
(12:30 PM) flew 12 feet out of her straight course to pass near one of
these shrubs. Two others were seen hovering around Wild Lime at 12:45
PM and 1 PM. Since Wild Lime flowers are small and inconspicuously
located in the axils, I assumed that the ponceanus females were seeking
nectar. But none of these shrubs proved to be in bloom.

On June 5 at 1 PM I followed a female as she fluttered through an
overgrown trail in a way that I had learned to associate with oviposition.
With wings vibrating closely together over her back, she perched for an
instant on a young shoot of Wild Lime at a height of 6 feet and laid an
egg on the leaf tip. I took this egg home to New York with me. The
larva which it produced is illustrated in the figures. It is also the basis
of some of the observations recounted below.

As will be apparent, this unexpected use of Wild Lime as a larval
foodplant offers a clue to the way in which ponceanus is adapted to its
habitat. Both Torchwood (the previously-recorded larval foodplant )
and Wild Lime are pioneering shrubs in whose shade sprout other hard-

130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

wood seedlings which can eventually grow up to hammock. Therefore
both grow abundantly at the edges of advancing hammocks but are
scarcer within the hammock itself, where they must struggle for light.
Wild Lime seems to need more sunlight than Torchwood. It often grows
completely exposed among low herbs, and conversely it is scarce in even
fairly open hammock. At the borders of hammocks both plants are usu-
ally densely-foliated shrubs up to 8 feet tall, but inside hammocks they
are slender trees over 15 feet tall, with fewer leaves. Presumably this
change to an arboreal habit is partly an adaptation to keep them from
having to struggle against their own shade in addition to the shade of
overtopping vegetation.

Most of the Torchwood and Wild Lime and the ponceanus associated
with them occurred in moderately shady hammock on rather bare lime-
stone pitted with sinkholes. Torchwood but not Wild Lime was nearly as
abundant in a still shadier habitat, proliferating on rich humus, but only
one ponceanus was seen there. This was a female found at 12:30 PM
flying higher than usual at about 15 feet within diffused sunlight just
below the forest canopy. In the shadiest place of all, a dense mature
hammock into which little light penetrated, no Torchwood and no
ponceanus were seen.

In contrast to the preference for partial shade shown by female pon-
ceanus, three female cresphontes were seen laying eggs on Wild Lime
growing on a similar limestone formation (11 AM in October, 1968).
Each cresphontes was completely exposed in bright sunlight, ovipositing
at a height of about 5 feet.

Eggs and Larvae. After learning to recognize ponceanus females in
search of the larval foodplants I was able to find eggs and larvae. It 11
AM on June 5, I followed a female to a spindly Torchwood tree growing
in the shade. She laid an egg on one of its new leaves. There were two
other eggs and eight larvae in various stages of development, even two
larvae in their third instar, on this one tree. On Torchwood trees in
similar situations I found other eggs and larvae but fewer of them. Eggs
were always on top of young leaves (but not always the youngest leaves )
and once on the rachis. Larvae were invariably on top of the youngest
and tenderest leaves, the new growth brought out by the spring rains.
Iiven the two third-instar larvae were feeding on young shoots and re-
fused older growth when it was offered.

No larvae were found on the few Wild Lime trees within the hammock,
nor on the numerous Torchwood and Wild Lime shrubs exposed to direct

unlight along the edges of hammocks. Three Papilio eggs were found

nlight at a heisht of 3 feet onaT h d 5
nt Or 3 teet on a Torchwood shrub growing half ex-

131

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VOLUME 25, NUMBER 2 133

posed along a trail. They did not produce detectable embryos. No
ponceanus eggs or larvae were found lower than 3 feet or higher than 7
feet from the ground.

First- and third-instar larvae which had been feeding on Torchwood
unhesitatingly accepted the youngest leaves of Wild Lime, and so did my
newly-hatched larva which had not yet eaten. This larva was later trans-
ferred to potted Torchwood which I had brought back to New York with
it. When the larva was in its second instar I ran out of yeung Torchwood
shoots and had to give it slightly older leaves. It refused them. Think-
ing that perhaps the waxy integument of an older leaf made it too tough
for food, I scraped the leaf edges with a razor blade, but the larva still
would not eat. Finally I gave this larva young shoots of Prickly Ash,
Zanthoxylum americanum Mill. from New Jersey. Prickly Ash, the pre-
ferred larval foodplant of cresphontes in the Northeast, grows in rocky
places (limestone ledges in New Jersey, sandstone outcroppings in IIli-
nois) much as the related ponceanus hostplants occur on well-drained
limestone in Florida.

Prickly Ash lacks the waxy covering of its subtropical relatives. Even
its mature leaves are tender. However, its youngest leaves are thick and
hairy, unlike the young shoots of Torchwood and Wild Lime. The
hungry ponceanus larva nibbled at the youngest growth of Prickly Ash
but seemed to have trouble eating the thick edges of these leaves despite
their tenderness. It reluctantly accepted leaves of intermediate age but
fed so seldom that its second instar lasted 7 days instead of 5. As the
larva matured it ate any Prickly Ash leaves offered, without apparent
preference for younger ones. It was reared to the pupal stage on this
substitute foodplant.

When kept at 80° F., about 5° lower than the average daytime tem-
perature of its natural habitat, the ponceanus larva was remarkably ir-
ritable. Confined in a vial, it would jerk its head backward if the vial
was jarred even slightly. If the vial was picked up, the larva shook its
head from side to side. Though newly-hatched wild larvae and my own
reared larva would protrude their white osmateria (producing an un-
pleasant smell) when I even touched the leaves on which they rested,
my larva as it matured into its fourth and fifth instars became reluctant
to do so even when prodded. At the same time it gradually abandoned
its habit of resting on top of leaves. Instead it clung to twigs below the
leaves when it was not actually feeding. Nearly mature cresphontes lar-
vae found on Prickly Ash in northern Illinois, rested in a similar position.

Grimshawe noted the curious thirst of ponceanus larvae in all instars.

134 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

In Florida, when I temporarily confined young wild larvae at 85° F. with
nearly saturated humidity, I never saw them drink. And when I reared
my larva in New York under similar conditions it apparently did not
drink during its earlier instars. But when in its fifth instar it was trans-
ferred to a roomy plastic box and kept at 80° F. with relative humidity
which varied from 60% to 70%, it would seek out and quickly drink
water droplets which I had scattered to keep the foodplant fresh.

Summary and Discussion

Papilio aristodemus ponceanus is found most often in old disturbed
places which are growing up to hammock. Feeding of larvae and adults,
courtship and oviposition occur in these second-growth hammocks, in-
dicating that ponceanus is a forest insect. The sympatric P. cresphontes
cresphontes feeds and reproduces in open places.

Ponceanus seems highly sensitive to moisture. It may emerge in num-
bers only after heavy or prolonged rains have ended the dry season and
brought out new growth on its host plants. A twenty-year summary of
South Florida weather records (Butson 1962) showed little change in
relative humidity between dry and rainy seasons. In the shelter of ham-
mocks, where there is usually no noticeable wind, the relative humidity
would be even more stable. However, Butson pointed out that the first
month of a rainy season usually has double the rainfall of the preceding
dry month. The beginning of the rainy season in South Florida varies
from late April to late June, the same months as the flight period of
ponceanus.

The ability of pupae to remain dormant for longer than a year as noted
by Grimshawe, may be a mechanism to synchronize adult emergence
with the beginning of the rainy season which in turn produces abundant
new growth for the larvae. In an exceptionally dry spring, emergence
could be delayed or suppressed. It is possible that pupae respond to
light intensity as well, for ponceanus does not emerge throughout the
rainy season as Antillean aristodemus populations apparently do.

The habitat of ponceanus is more or less continuous except where man
has destroyed it. But feeding adults disperse widely. Their distribution
seems related to the freshness of Guava blossoms, the preferred adult
foodplant within hammocks at the time of my observations. Adults would
not be found at the same Guava patches for more than a day. However,
one patch of Cheese Shrub blooming in the open attracted a male a day
lor three days. The short duration of Guava blossoms may aid the repro-
ductive dispersal of ponceanus.

In contrast with the fluid distribution of feeding adults, the eggs and

VoLUME 25, NuMBER 2 1335)

larvae were distributed very narrowly. The single Torchwood tree which
yielded eight larvae in various stages of development seems to indicate
that it or its situation was especially attractive to several ovipositing fe-
males. This contention seems supported by the fact that other Torchwood
trees of the same size growing in similar conditions, generally had larvae
on them.

Young larvae will accept other foodplants rather than eat any but the
tenderest leaves of the “right” foodplant. They will feed on at least two
other Rutaceous plants besides Torchwood, the primary hostplant. Ovi-
position on one of these, Wild Lime, occurs in nature. Of course ready
acceptance of Wild Lime does not necessarily mean that most larvae will
thrive on this diet and mature into adults capable of reproducing. Never-
theless, the assured behavior of the female which oviposited on a Wild
Lime tree growing in the shade, as compared with the hesitation of the
three females seen around Wild Lime shrubs growing in the open, sug-
gests that specific habitat might be a stronger stimulus to oviposition than
specific foodplant. The fact that ponceanus females are only stragglers
in open places and that Wild Lime is only a straggler in shady places
makes it unlikely that oviposition on this secondary hostplant is frequent.

It is remarkable that the shady habitat of ponceanus is not the most
favorable place for its primary hostplant either. As the price of reproduc-
tive isolation ponceanus lives cut of step with both Wild Lime and
Torchwood. Both are constantly being shaded out of hammocks, each
at its own rate, by normal plant succession.

The mechanisms which maintain environmental separation between
ponceanus and cresphontes are not fully understood. An initial hurdle of
reproductive isolation may be environmental separation of virgin females
which are feeding. Fertilized female ponceanus may have a genetically
controlled preference for partial shade when ovipositing. As for males,
there seems to be a tension zone at the edges of hammocks, characterized
by occasional combat between cruising males of ponceanus and cre-
sphontes.

In the mottled light of its habitat ponceanus may benefit from a pat-
tern resemblance to Heliconius charithonius tuckeri Comstock & Brown.
I was sometimes unsure whether a yellow-streaked butterfly vanishing
into the edge of a hammock was a rare swallowtail or merely a common
heliconian. When I saw such a butterfly from behind, foreshortening in-
creased the resemblance.

It is well known that the wing patterns of H. c. tuckeri and P. cre-
sphontes are “hard edged” while that of P. a. ponceanus is suffused. Pat-
tern suffusion is a reputed characteristic of tropical mimetic butterflies

136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(Corbet & Pendlebury 1956), perhaps indicating that the need for close
resemblance to a model is reduced by the preference for shady habitats
which such mimics often show. I have already mentioned the “agreeable”
scent of ponceanus males and the “disagreeable” scent of a male cre-
sphontes from the same place. Further investigation could determine
whether it is more than coincidence that some cresphontes hostplants
belonging to the genera Ruta, Dictamnus and Ptelea have been respon-
sible for photosensitive dermatitis in mammals (Muenscher 1951). My
few dried Torchwood specimens have more and larger resin dots on young
leaves from a completely exposed plant than on young leaves from a plant
which grew in partial shade. It is conceivable not only that South Florida
cresphontes might be protected by an irritant substance in some native
Rutaceae, but also that shade-grown Torchwood and Wild Lime might be
deficient in that irritant.

Putting conjecture aside, there is reason to believe that Papilio aristo-
demus ponceanus must continuously colonize new places partially de-
forested by storm or fire, which temporarily are in a state of regrowth
suited to its habits. The extremes of heedless real-estate development
and over-protective maintenance of mature hammocks both threaten to
disrupt the natural cycle of renewal on which its survival depends.

Acknowledgments

I am indebted to Mr. George N. Avery of the Fairchild Tropical Gar-
dens, Miami, Florida, for identifying Cheese Shrub, to Dr. C. F. dos
Passos of Mendham, New Jersey, for editorial help with part of the
manuscript, to Mr. Winthrop Edey of New York City for the photographs,
and to Dr. A. B. Klots of the American Museum of Natural History for
information which made my observations possible.

Literature Cited

Burson, K. 1962. Climate of Florida. In Climatography of the United States No.
60-8. Revised edition. Washington.

Corser, A. S. anp H. M. PENDLEBURY. 1956. The butterflies of the Malay
Peninsula. Second edition revised. Edinburgh.

GRIMSHAWE, F. M. 1940. Place of sorrow. Nature Magazine 33: 565-567, 611.

Henperson, W. F. 1945a. Papilio aristodemus ponceana. Ent. News 56: 29-32.

1945b. Additional notes on Papilio ponceana. Ent. News 56: 187-188.

ere 1946. Papilio aristodemus ponceana Schaus notes. Ent. News 57: 100-

Ml.
KimBaLL, C, P. 1965. The lepidoptera of Florida. Gainesville.
Kiots, A. B. 1951. A field guide to the butterflies. Boston.

Mr ee ni C. 1951. Poisonous plants of the United States. Revised edition.
YCW OrK,

VoLuME 25, NuMBER 2 137

NOTES ON SOME SOUTH FLORIDA LEPIDOPTERA

FRANK RUTKOWSKI
153 Centre Street, New York, N.Y.

The following observations from the Florida Keys are additions to in-
formation published by Kimball (1965). These observations may be of
interest because they represent apparently new records of habits, larval
foodplant or extension of known range. Nomenclature and checklist num-
bers follow dos Passos (1964) for butterflies and McDunnough (1938)
for moths. Plant names are taken mostly from Small (1933). I am in-
debted to Mr. George Avery of the Fairchild Tropical Gardens in Miami,
Florida, who helped me in various ways and was also kind enough to
identify two of the more difficult plants.

29. Lerodea eufala (Edwards). Extension of range. A few of these
skippers were taken in December on Big Pine Key, visiting blossoms of
Croton linearis Jacq. in the shadier parts of pine woods.

281. Ascia monuste phileta (Fabricius). During a migration of this
species on Key Largo in June, a pair was found in copula within shady
hammock at 1:15 PM on a sunny day, the dark female flying.

340. Lephelisca virginiensis (Guérin-Méneville). Extension of range.
A colony was found in August on Big Pine Key, in rocky pinelands imme-
diately adjacent to marshes.

380. Strymon martialis (Herrich-Schaffer). This very local species
seems to prefer the blossoms of Bay Cedar (Suriana maritima L.) when
available, to the flowers which it usually visits at other times.

45la. Hemiargus ammon bethunebakeri Comstock & Huntington. New
larval foodplant. At 10 AM in early June a female laid an egg just above
a lateral bud on Snowberry, Chiococca alba (1L.) Hitche., growing along
a shady trail on Key Largo.

486c. Anaea aidea floridalis Johnson & Comstock. A butterfly trapped
in a spider web was tasted and immediately discarded by the spider.

524b. Metamorpha stelenes biplagiata (¥Frihstorfer). Extension of
range. In late October a colony was found on Big Pine Key, associated
with Blechum pyramidatum (Lam.) Urban, which is probably the larval
foodplant. The butterflies are very fond of the blossoms of Wild Lantana,
Lantana involucrata L. When at rest with wings closed, they are hard
to tell from the curled and blotchy leaves of Jamaica Dogwood, Ichthyo-
menthia piscipula (L.), which start to turn brown and fall at this time of
year. One female apparently had been attracted to light, for early on

138 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

a rainy morning she was found among moths in the grass at the base of
a streetlamp.

531. Junonia coenia (Hubner). This species and Junonia evarete
zonalis Felder & Felder interbreed on Big Pine Key. J. e. zonalis is com-
mon along mudflats while J. coenia seems to prefer damp places on
higher ground. The roads and fire lanes which have been opened on
Big Pine Key may help to break down reproductive isolation of these two
populations. Copulating pairs representing various intergradations were
taken around 5 PM in December, when the sun was quite low. The fact
that mating took place late in the day when light had shifted from the
violet to the red end of the spectrum might have minimized any im-
portance that pattern or color differences could have in courtship. If
this interbreeding really represents secondary intergradation, it is to be
expected that the two populations might formerly have diverged even less
in courtship ritual than in appearance. The instance of abortive courtship
listed below under Euptoieta suggests that in bright sunlight any rusty
orange butterfly resting on the ground in a likely habitat might momen-
tarily arouse the interest of a male J. e. zonalis.

624. Euptoieta claudia (Cramer). Extension of range. Occasional but
fresh specimens were taken on Big Pine Key in May, August and October.
They were found in both very wet and very dry habitats. One female
briefly responded (spreading out of wings and raising of abdomen) to a
hovering Junonia evarete zonalis.

628d. Dryas julia cillene (Cramer). Extension of range. A battered
female was taken in August on Big Pine Key along the edge of dry ham-
mock on an outcropping of Key Largo Limestone.

818d. Automeris io lilith Stkr. New larval foodplant. Larvae were
found on Croton linearis on Big Pine Key in December.

560a. Eacles imperialis didyma Beauv. Extension of range. A badly
smashed specimen was found among other moths at the base of a street-
lamp on Big Pine Key in October.

570. Lymire edwardsii Grt. Numerous cocoons were found at the base
of a Sapodilla, Sapota Achras Mill., trunk on Big Pine Key in August.

1954. Xanthopastis timais Cram. Third and fourth instar larvae found
skeletonizing the leaves of Hymenocallis keyensis Small on Big Pine Key
in October, freely accepted Iceberg Lettuce as a substitute foodplant in
New York and were reared to maturity on it.

3516a. Composia fidelissima vagrans Bates. One of these moths was
removed unharmed from a spider web on Big Pine Key in December.
[ nlike netted specimens, it responded to being handled by squeaking and
excluding a drop of yellowish froth at each side of the prothorax. This

VoLuME 25, NuMBER 2 ifs9

froth had a very faint but disagreeable taste which can be compared to
the smell of a “sour” dishcloth.

5221. Sphacelodes vulneraria (Hbn.). Extension of range. A pair was
caught at light on Big Pine Key in October.

Literature Cited

pos Passos, C. F. 1964. A synonymic list of the nearctic Rhopalocera. Mem.
Lepid. Soc., No. 1.

KimBaLL, C. P. 1965. Lepidoptera of Florida. Div. of Plant Industry, Fla. Dept.
of Ag., Gainesville.

McDunnouwcu, J. 1938. Checklist of the Lepidoptera of Canada and the United
States of America. Part 1. Macrolepidoptera. Mem. Southern Calif. Acad. Sci.
olor,

SMALL, J. K. 1933. Manual of the southeastern flora. Univ. of North Carolina
Press, Chapel Hill.

NYMPHALIDAE OF WISCONSIN

Davin WM. SCHWEHR
University of Wisconsin, Madison

Fernekes (1909) and Muttkowski (1907) compiled a list of over 1000
species of Lepidoptera, including Nymphalidae, found in the Milwaukee
County area. The list was by no means a complete survey of the Lepidop-
tera of Wisconsin, nor was it entirely accurate for the Milwaukee area.
Lack of extensive collecting, deficiency of accurate records, and absence
of recent distribution studies have resulted in only generalized records on
the distribution and range of Nymphalidae in Wisconsin.

Thus, during the summer of 1966 and continuing into the spring of
1968, a survey was conducted in an attempt to establish the distributive
range of Nymphalidae in Wisconsin. Literature sources provided a pre-
liminary list of Wisconsin Nymphalidae. Specimen records were then
obtained from the collections of the University of Wisconsin, Northern
Michigan University, University of Minnesota, Concordia College, Wis-
consin State University at Stevens Point, and from the personal collection
records of J. A. Ebner, Wm. E. Sieker, Stephen C. Kleene, and Kurt
Johnson, as well as from my own collection.

Twenty-four species of Nymphalidae were obtained through personal
collecting in various parts of Wisconsin, ranging from Bayfield County in
the north to Dane County in the south. Polygonia faunus, Polygonia
progne, Phyciodes batesii, Phyciodes gorgone, and Melitaea harrisii were
added through reference to the collection of the Russel Laboratory, at
the University of Wisconsin. Collection records of Agraulis vanillae and

140 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Boloria freija were received from Wm. E. Sieker, Madison, and Kurt
Johnson, Stevens Point. Records of Anaea andria and Nymphalis califor-
nica have also been reported from the State by J. A. Ebner, West Allis.

Thirty-three species representing eight subfamilies and fifteen genera
have been collected within the boundaries of Wisconsin. Polygonia
gracilis and Polygonia satyrus may also occur within the State. A speci-
men of the latter, in the collection at Northern Michigan University, was
taken in northern Wisconsin. However, its presence has not been estab-
lished due to lack of collecting in that part of the State.

The county records for individual species are available to the interested
reader. The following are the generalized ranges of Wisconsin Nym-
phalidae:

Charaxinae
Anaea andria Scudder. At present recorded only from Racine Co., but
may inhabit the southern third of the State.

Apaturinae
Asterocampa celtis (Boisduval & Leconte). Probably present in the en-
tire State, but not yet recorded from the northwestern counties.
Asterocampa clyton (Boisduval & Leconte). Inhabits the southern two-
thirds of the State.
Limenitidinae
Limenitis (Limenitis) arthemis (Drury). Occurs through most of Wis-
consin, but its existence is doubtful in the southern extremities.
Limenitis (Limenitis) astyanax (Fabricius). Is found in the southern
half of Wisconsin.
Limemitis (Limenitis) archippus (Cramer). Has a state-wide range.

Vanessinae
Vanessa atalanta (Linnaeus). Has a state-wide range.
Vanessa virginiensis (Drury). Has a state-wide range.
Vanessa cardui (Linnaeus). Has a state-wide range.
Junonia coenia (Hubner). Is scattered throughout Wisconsin.

Nymphalinae
Nymphalis vau-album (Denis & Schiffermuller). Has a state-wide range,
however the species is most frequently collected in northern Wisconsin.
Nymphalis californica (Boisduval). Has only been collected once in
Wisconsin, and that was in Door Co.
Nymphalis milberti (Godart). Has a state-wide range.
Nymphalis antiopa (Linnaeus). Has a state-wide range.
Polygonia interrogationis (Linnaeus). Has a state-wide range.

VoLUME 25, NuMBER 2 141

Polygonia comma (Harris). Has a state-wide range.

Polygonia faunus (Edwards). Has been collected in the northern coun-
ties of Door, Chippewa, and Marinette.

Polygonia progne (Cramer). Ranges throughout Wisconsin; however
the species is more frequently collected in the northern territories.

Melitaeinae

Phyciodes (Phyciodes) tharos (Drury). Has a state-wide range.

Phyciodes (Phyciodes) batesii (Reakirt). Has been taken only in the
northeastern counties of Bayfield, Brown, and Marinette.

Phyciodes (Phyciodes) gorgone (Hiibner ). Is scattered throughout Wis-
consin.

Melitaea ( Microtia) harrisii Scudder. Inhabits portions of eastern Wis-
consin, being most frequently collected in the northern third of the
State.

Melitaea ( Microtia) nycteis Doubleday. Has a state-wide range.

Euphydryas phaeton (Drury). Has a state-wide range.

Argynninae

Boloria (Clossiana) selene (Denis & Schiffermuller). Has a state-wide
range.

Boloria (Clossiana) toddi (Holland). Has a state-wide range.

Boloria (Clossiana) freija (Thunberg). Has only been collected in Bay-
field, Oneida, and Portage counties, but may later be found to range
throughout the northern counties.

Speyeria (Speyeria) idalia (Drury). Distributed throughout southern
Wisconsin.

Speyeria (Speyeria) atlantis (Edwards). Distributed through the north-
ern third of Wisconsin.

Speyeria (Semnopsyche ) cybele (Fabricius). Has a state-wide range.

Speyeria (Semnopsyche) aphrodite (Fabricius). Has a state-wide range.

Euptoieta claudia (Cramer). Is scattered throughout Wisconsin.

Heliconiinae
Agraulis vanillae (Linnaeus). Has only been collected in Grant and
Portage counties, but may later be found to range throughout the
southern parts of Wisconsin.

Acknowledgments
I am grateful to Dr. R. D. Shenefelt and L. J. Bayer, the University
of Wisconsin; Gordon D. Gill, Northern Michigan University; Dr. Ke
Chung Kim, the University of Minnesota; George Senechal, Concordia
College; Kurt Johnson, Wisconsin State University at Stevens Point; J. A.

142 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Ebner, West Allis, Wisconsin; and Wm. E. Sieker, Madison, Wisconsin,
for their cooperation in providing collection records.

Sincere appreciation is also extended to both the Wisconsin Junior
Academy of Science, for its grant for work on insects in 1967-68, and Mr.
Leroy Lee, Biology teacher at James Madison Memorial High School, for
his assistance and guidance throughout the research.

Literature Cited

pos Passos, Cyrit F. 1964. A Synonymic List of the Nearctic Rhopalocera. Lep.
Soc. Mem. 1.

ExrucuH, Paut R. anp ANNA H. ExneuicH. 1961. How to Know the Butterflies.
Brown Pub. Co., Dubuque.

FERNEKES, VALENTINE. 1906. List of Lepidoptera occuring in Milwaukee County,
Bull. Wisconsin Nat. Hist. Soc. 4.

Kiors, ALEXANDER B. 1964. A Field Guide to the Butterflies of North America,
East of the Great Plains. Houghton-Mifflin, Cambridge.

Murrkowskti, Ricuarp A. 1907. Additions to the Lepidopterous Fauna of Mil-
waukee County. Bull. Wisconsin Nat. Hist. Soc. 5.

A NEW NAME FOR PAPILIO ZETES WESTWOOD, 1847,
NEC LINNAEUS, 1758 (PAPILIONIDAE)

EUGENE MUNROE
Entomology Research Institute, Canada Department of Agriculture, Ottawa

J. O. Westwood (1847, p. 36, pl. 3, figs. 1, 1*) described Papilio zetes
from St. Domingo. This was recognized as a valid species of Papilio by
subsequent authors, e.g., Rothschild and Jordan (1906, p. 515), but was
transferred by Munroe (1961, pp. 47, 51) to Battus Scopoli. D’ Almeida
(1966, p. 116) followed this placement.

Unfortunately Papilio zetes Westwood is a junior primary homonym of
Papilio zetes Linnaeus (1758, p. 487), now placed in Acraea Fabricius,
and it has no junior synonyms. Therefore, under Article 60(b) of the
International Code of Zoological Nomenclature, the former name requires
replacement. I propose Papilio zetides, nom. noy., to replace Papilio
zetes Westwood. Papilio zetides should be transferred to Battus, becom-
in Battus zetides (Munroe), comb. nov.

Literature Cited

D’AtMeipA, R. Ferrera, 1966. Catalogo dos Papilionidae Americanos. Soc.
Brasileira de Entomologia, S40 Paulo.

LINNAEus, C. 1758. Systema Naturae, ed. X.

Munroe, E. 1961. The classification of the Papilionidae. Mem. Ent. Soc. Canada
Ag

Roruscuitp, W., AND K. Jorpan. 1906. A revision of the American Papilios.
Nov. Zool. 13: 411-752, pl. 4-9.

Wesrwoop, J. O. 1847. Description of two new exotic species of the genus
Papilio, Trans. Ent. Soc, London 5: 36-37, jolly,

VoLUME 25, NuMBER 2 143

NOTES ON THE OCCURRENCE OF TWO RARE LEPIDOPTERA
IN SOUTH CAROLINA

In late May and early June of 1970, I discovered a colony of Satyrium kingi (Klots
and Clench) in Dorchester County at the county line on highway 642 where Dor-
chester and Charleston counties meet. As far as I have been able to determine this
is the first Dorchester County records, and possibly the first record for the state in
the Coastal Plain.

In July, a colony of Euphyes bimacula (G. & R.) was found just east of Summer-
ville, S. C. in Berkely County near the junction of U. S. Al7 and I-26, and to the
northwest of this junction. According to Klots (1951, A Field Guide to the Butter-
flies ) this is well south of its supposed range.

RONALD R. GATRELLE, 35 Reddin Rd., Apt. 1, Charleston, South Carolina.

_ VAN SOMEREN BUTTERFLY COLLECTION
TO THE AMERICAN MUSEUM OF NATURAL HISTORY

This magnificent collection of the butterflies of East Africa, mostly from Kenya,
Uganda, and north Tanganyika, plus the adjacent areas of southern Ethiopia,
Somali, and eastern Congo, has been donated to the American Museum of Natural
History. It represents over 60 years of work by Dr. V. G. L. van Someren, with
the specimens being obtained both by catching and by rearing; it is one of the largest
and most complete collections of butterflies from the above area. The collection
consists of 22,931 specimens; of this total, 18,497 are butterflies; there are 258
slides of genitalia, and 4,176 specimens of early stage material, particularly of
Charaxes. As far as I know, this is the largest single collection of African butterflies
to come to the United States at one time.

The specimens of this collection have been studied and identified by many special-
ists and have been included in their revisionary studies on African butterflies. These
include W. H. Evans’ “A Catalogue of the African Hesperiidae” (1937), H. Stempf-
fers “The Genera of the African Lycaenidae” (1967), and Dr. van Someren’s own
“Revisional Notes on African Charaxes” (1963-1969, with more parts to be pub-
lished ), just to mention a few. Dr. van Someren has reared many species of Charaxes
and has made a sizable collection of cast larval head capsules and of pupae; this
valuable material came with the collection.

No holotypes or allotypes are included with the collection. Dr. van Someren has
deposited his type specimens, as well as a portion of his butterfly collection, in the
entomological section of the British Museum (Natural History).

FREDERICK H. RinpcE, Dept. of Entomology, American Museum of Natural History,
New York.

A MELANIC ABERRATION OF PHYCIODES THAROS
(NYMPHALIDAE )

A striking female aberration of Phyciodes tharos (Drury) emerged 29 May 1969
from a laboratory brood reared from a wild-inseminated female collected four miles
east of Cedar Key, Levy Co., Florida, 29 March 1969. All rearing was done using
a 16-hour daily photophase at 27° C and with Aster ericoides L. as the larval food-
plant.

A comparison of the melanic female with a normal sibling female shows that all
of the black pattern elements of both the dorsal and ventral wing surfaces have
spread into the fulvous areas between them, leaving light pigmentation only in the
central parts of normally fulvous areas. The ventral “pearl crescent” near the anal
margin of the hind wing has been completely covered by dark pigmentation, but it

144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

can be seen through the black scales and is otherwise normal. The coloration of the
body and of the appendages has remained unaffected.

There is a melanic P. tharos female labelled “Norwich, Mass./30 July 1955/leg.
M. Cady” in the Peabody Museum at Yale, but in this specimen the wing markings
have become completely blurred and smudged. On the dorsal side there is exten-
sive fulvous at the wing bases with the rest of the wing surface black. Ventrally
only two of the forewing black markings appear on a clear yellow ground, and the
hind wing is cream with a large central brown patch. The two melanic specimens
are very different in the way that the wing patterns have been affected, and they
may be the result of quite different effects (for example, environmental vs. genetic).

The Florida individual was the last to emerge in a brood of 158 individuals
(876 6, 6192 9), pupating and emerging several days after the last of its siblings.
The melanism is thus correlated with a significant slowing in the rate of develop-
ment, due probably either to a direct effect on the developmental rate or to a gen-
eral lessening of vigor.

It is interesting to speculate on the possible inheritance of the form. The melanic
was mated to a non-melanic sibling and produced a brood of about 40 adults, all
of non-melanic appearance. Embryo mortality was high but normal for an F: of a
sibling mating in this species; larval and pupal mortality were negligible. If the
melanic form is genetic in origin, it is probably recessive.

Cuarces G. Oxiver, Hope Dept. of Entomology, Oxford University, Oxford, En-
gland.

A SPECIALIZED CASE OF COMMUNAL ROOSTING IN
PIERIS RAPAE (PIERIDAE)

As a possible parallel to the report by Clench (1970, J. Lepid. Soc. 24: 117-120)
it seems worthwhile to record my observations on a roosting aggregation of Pieris
rapae (L.) in a garden at 2 Gulfview Rd., Blackwood, South Australia, in the foot-
hills of the Mt. Lofty Range south of Adelaide, at an elevation of 800 feet. I lived
at this address for five and one half years (1965-1970), during which time the fol-
lowing was observed.

P. rapae is abundant and multiple-brooded in this locality; the larvae feed upon
a luxuriant patch of nasturtium (Tropaeolum) in the garden being described. Adults
are on the wing from early spring (Sept.) to autumn (Apr.-May), reaching a peak
during the summer period (late Nov. to early March). In this garden grows a small
tree (height approximately 10 feet; shape roughly pyramidal; foliage cover fairly
open) of Pittosporum undulatum Vent. var. variegatum (PITTOSPORACEAE).
The important feature of this tree, with respect to the account that follows, is the
coloration of its leaves. The leaves of this smaller-growing garden variety of P.
undulatum are a pale greenish-white with contrasting white margins. The overall
effect of the tree color is whitish-green; seen in the warm yellowish rays of
late afternoon sunshine, just prior to sunset, these leaves light up with a faintly yel-
lowish or cream-green tinge which is very close to the shade on the visible areas
of the undersides of P. rapae wings when the butterflies are in their normal resting
position with the wings closed over the dorsum. This P. undulatum tree was growing
among other nearby trees, shrubs, and vines, but was in an opening where it received
direct sun for most of the day. During sunny summer days, adults of P. rapae flew
through this garden by the dozens, often five or more being visible at one time within
the boundaries of the garden. Many of them would fly over and around this tree
during the midday hours, even then showing somewhat more attraction to it than

VoLUME 25, NuMBER 2 AS

to most other vegetation in the garden. Their primary attention, however, was cen-
tered around the extensive nearby Tropaeolum patch during this part of the day.

As late afternoon approached, on most sunny days, many individuals would be
seen circling, alighting upon, or flying in and out of the small Pittosporum tree.
Their concentration around this particular tree would become quite noticeable,
although a few would also be seen flying around other plants in the same garden,
preparatory to settling for the night. As the sun came closer to the horizon, more
and more individuals settled in this tree. At first there would be much alighting
and. taking off again, and fluttering slowly about, before final settling took place.

Once at rest among these whitish-green leaves, the butterflies were very hard to
see at a passing glance, although careful scrutiny would show up some of them. Num-
bers settling into this tree would often approach 10 or more individuals by sunset,
whereas other nearby vegetation would only attract the odd individual. A person
passing the Pittosporum tree in late afternoon, at the right time, would often cause
a small “cloud” of P. rapae to burst from the tree as he passed by; later (early eve-
ning) they would not arise from the tree unless it was knocked, or a very sudden
movement was made at close range. During the period of “settling in” I noticed
that most of them landed on the western (sunny) side of the tree, and among its
upper branches, wherever the last weak rays of sunshine remained longest.

This case of “communal” roosting is probably more or less explained in the last
paragraph of Clench’s discussion. It seems likely that the distinctive foliage colora-
tion of this particular variety of P. wndulatum, plus its favorable location with re-
spect to the late rays of sunlight, combined to provide a special attraction for P.
rapae under the influence of late afternoon sunlight. In most other circumstances,
I strongly suspect that this pierid would show little or no communal roosting be-
havior. At best they might show only a slight attraction for one type of roosting
situation over some others, but they would probably not form any noticeable aggrega-
tions on any single plant.

No marking of individuals was undertaken during these summers, so I cannot say
whether the same individuals came to roost in this same tree night after night, or
if there was any tendency to use certain leaves or stems repeatedly in preference to
others on the same tree. I would guess, however, that a fairly large percentage of
different or new individuals formed the aggregation in this tree each evening. Prob-
ably the turn-over was very high each day, as the butterflies wandered through this
residential area, from garden to garden. Those finding themselves (by late after-
noon) in the garden described, would naturally gravitate toward the most attractive
roosting place which, in this instance, happened to be the small creamy-green
Pittosporum tree.

Noet McFar.anp, 129 Gloucester Ave., Belair, South Australia.

NOTES ON THE CONFUSION BETWEEN LETHE CREOLA AND
LETHE PORTLANDIA (SATYRIDAE)

There has been confusion between Lethe creola (Skinner) and Lethe portlandia
portlandia (Fab.) in not only private collections but also in institutions and in litera-
ture.

This confusion is evidenced by the fact that the female of the type series of creola
from the Skinner collection, which is now in the Carnegie Museum, was found to be
a female of L. portlandia by Gillham and Ehrlich. Without going further into

146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

this aspect of the matter I will simply refer the reader to the 1970 article by Roderick
R. Irwin the (Jour. Lepid. Soc. 24: 143-151).

While collecting these two species in South Carolina (1970), I found a constant
pattern in the flight habits of the two species. It became evident why there has
been so much confusion between them. Confusion exists in regard to the females;
the males of the two species are easily distinguished. Many of the earlier collectors,
especially in Louisiana where the type and allotypes of creola came from, must have
encountered the same situation which I did. The habitat of creola is often an in-
accessible area of swampy, bushy, cain-filled undergrowth. Naturally one would
tend to collect in the more open areas in this type of terrain. In the open areas
where collecting is more easily done you will find a preponderance of male creola
and female portlandia; both portlandia males and creola females are rare.

The following records were taken from 9 April to 18 Oct. 1970. Of 40 female
portlandia examined, 29 were collected in more open areas, such as along paths; 11
were taken in denser areas, e.g. 20 feet or more away from clearings. Only two male
portlandia were taken in open spaces whereas ten were caught in the denser areas.

Of 24 male creola caught, 15 were found in the open areas, nine in dense areas. Of
seven female creola found, five were in dense areas, only 2 in more open terrain.

The majority of specimens were released. Due to the difficulty in moving around
in the denser areas many specimens seen there escaped capture. The tendency of both
species to occur in different areas was not affected with regard to the time of year
but specimens were more difficult to capture in the fall.

RONALD R. GATRELLE, 35 Reddin Rd., Apt. No. 1, Charleston, South Carolina.

REMARKS ON “DISTRIBUTIONAL NOTES ON THE GENUS MESTRA
(NYMPHALIDAE) IN NORTH AMERICA”

I wonder if Mr. Masters is not attaching undue importance to the occurrence of
Tragia in his paper entitled as above (1970, Journal Lepidopterists’ Society, 24: 203).

Five species of East African Eurytelinae have their food-plants recorded, viz.
Byblia acheloia Wllgrn. and B. ilithyia Drury feeding on Tragia brevipes and Dale-
champia hildebrandti, Eurytela hiarbas Drury and E. dryope Cr. feeding on Dale-
champia hildebrandti and Ricinus communis and Neptidopsis fulgurata Bsd. re-
corded from Dalechampia hildebrandti only. The Indian Ergolis ariadne Johan. feeds
on two species of Tragia, whilst E. merione Cr. feeds on Castor (Ricinus communis).

I cannot help feeling that Mestra amymome may also have one or more alternative
foodplants.

D. G. SEvAstroputo, P. O. Box 5026, Mombasa, Kenya.

DRAGONFLY ATTACKS LIMENITIS DEFENDING ITS TERRITORY

On June 23, 1970, while collecting Limenitis archippus floridensis Strecker near
Folkston, Georgia, I observed a rather unusual sequence of events involving a male
Limenitis and a large dragonfly.

The Limenitis flew over a small shaded waterhole along Route 252. As I pursued
it, I observed the dragonfly dive at the Limenitis who evaded it and landed on a
cypress branch. After resting, the butterfly soared slowly over the open water. The
dragonfly swooped down and grasped the butterfly, then carried it to the water
where it was released.

The stunned butterfly fluttered weakly to a nearby branch, rested there a con-
siderable period of time flexing its wings frequently. The dragonfly soared past it
several times feigning attack each time the butterfly folded its wings. A final attack
by the dragonfly knocked the butterfly to the ground; it remained a few seconds

VoLUME 25, NuMBER 2 147

flexing its wings, then retreated into the densely wooded swamp with the dragonfly
in pursuit.

Dr. Clifford B. Knight states in Basic Concepts of Ecology, p. 157: “Dragonflies
will establish a linear territory along a stream or in the vicinity of a body of water
that they patrol and defend against invasion by other members of their species.”
Territoriality is normally intraspecific—could this unusual behavior suggest another
predator for the tasty Limenitis or the extension of territoriality to an interspecific
activity by the dragonfly?

During the past ten years, I have collected thousands of Limenitis. Always these
individuals were found in association with water, and in the South with the ever
present dragonfly which apparently shares its habitat. In most cases, this appears
to be a harmonious relationship.

THomas R. MANLEy, Bloomsburg State College, Bloomsburg, Pennsylvania.

THE OCCURRENCE OF VANESSA CARDUI IN MISSISSIPPI AND
TENNESSEE?

C. B. Williams (1970, Jour. Lepid. Soc. 24: 157) stated that V. cardui “is not
often recorded in the southeast, though if this is due to a real rarity or to lack of
interest is not certain.” Speaking of its occurrence in 1952, he wrote, “There are
however no records of abundance from Texas or from any of the Gulf States except
Mississippi, where it was said to have been ‘abundant’.” He concluded with a plea
for sharing one’s observations. I give here observations made in Mississippi and
Tennessee.

V. cardui was first recorded from Mississippi by Weed in 1894 as taken by him
in the northeastern part of the state during the three previous years. He reported
it rarer than virginiensis. Mather and Mather in 1958 reported having found it in
all months except January, May, June, and December. Records are now available for
May, June, and December, leaving only January without records. Localities were
known in nine counties in all sections of the state. It is probably their record of 1952
occurrence as “abundant” that is referred to by Williams. All data now available
to me for Mississippi occurrences are tabulated below in terms of number of recorded
occurrences per month. For the years not listed there are no recorded occurrences.

F M A M J J A S O N D
1947 — - - - _ — = 1 = - ~ il
1948 = = = = = = as 1 = a Es 1
1949 - = i = = _ ] 1 ] = = 4
1952 IL 3 iL 2 — 1 I 3 2 wks
1953 os 2) - - = > 1 - ~ IL _ 4
1954 = = Il = = ~ - = 2, 2 ~ 5
1957 - = = ~ - 2 3 1 I Ss = 7
1958 = = = I 2) 4 4 = - = = fil
1960 = a = il = = - = 2 = = 3
1965 = = — = - = 2 if = = = 3
19687 = _ 3 4 i = it il = 1 alll
1970? = = ~ _ I _ il 2 1 1 6

] 5 6 8 4 i. Sle! 9 10 il I Nea SS,

1 Contribution No. 190, Bureau of Entomology, Division of Plant Industry, Florida Department
of Agriculture and Consumer Services, Gainesville.
2 Includes data furnished by Mr. Charles T. Bryson, Mississippi State University.

148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Mr. Harry C. Monk, Nashville, Tenn. has tabulated his observations of the oc-
currence of V. cardui in that area (Davidson County) and has granted me permis-
sion to include them in this report. In the following tabulation of number of days
in any one month that V. cardui was observed, a day is counted regardless of the
number of individuals seen or the number of localities at which the species was
seen. Data through 1970 are included. He has one December record: 8 December
1952; this is the only year for which there is a December record for Mississippi.

APR MAY JUN JUL AUG . SER. OGi@Gr
1954 - ~ It 9 4 24 1 2 57

1955 ~ - - - - 3 3 - 6
1956 - - = _ 2 il - ~ 3
1957 - - - 9 11 19 10 1 50
1958 - - 3 13 10 iky/ 1 1 45
1959 - - iL i - 2 - - 4
1960 - - - + J 9 5 - 29
1961 - — — 2 I - - - 3
1962 ~ _ - - 3 + 2 3 12
1963 - - 2 2 a 6 9 il 27
1964 _ _ - - 2 2 Uh 1 12
1965 - - - if 6 6 2 — 15
1966 1 2 7 1 8 7h 12 2 40
1967 - - ~ - - - 5 - 5
1968 4+ 6 10 5 25 13 9 - 72
1969 - - - - - - 1 - 1
1970 - ~ I 2 4 5 2 i 15

5 8 25 AQ 94 118 85 12 396

BryAnr MATHER,’ 213 Mt. Salus Dr., Clinton, Mississippi.

3 Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services.

BOOK REVIEW

THE ENciisH LEPIDOPTERA OR THE AURELIAN’S POCKET COMPANION, by Moses
Harris. 1775, 66 pp. + i-xv, 1 coloured plate. Reprinted 1969 by E. W. Classey
Ltd., Hampton, Middlesex. Distributed in North America by Entomological Reprint
Specialists, P.O. Box 77971, Dockweiler Station, Los Angeles, Calif. Price $9.60 U.S.

An introductory section of the book is devoted to the collection, rearing and pres-
ervation of moths and butterflies. The remainder of the text is arranged in the
form of a table which lists for 415 species of British Lepidoptera: common names,
food plants, seasonal periods of pupation and emergence, wing expanses, habitats, and
diagnostic macular features. The last column in the table of data lists the Linnaean
names for each species and numbers them according to the tenth edition of the
Systema Naturae. A coloured frontispiece illustrates the numerial system of designat-
ing wing veins and cells. The book will be a particular interest to the bibliophile.

D. F. Harpwicx, Editor.

VOLUME 25, NUMBER 2 149

ADDITIONAL RECORDS OF PATRICIA DEMYLUS GEMELLUS
FOX (ITHOMIIDAE)

Patricia demylus gemellus Fox was described (1960, Jour. New York Ent. Soc. 68:
152-156) from only three known specimens (two in the British Museum and one
in the Museum of Comparative Zoology ), all males, and all from Bolivia.

I have obtained eleven additional specimens in consignments from Franz Stein-
bach of Cochabamba, Bolivia. The first eight of these (including four males and
two females from E] Limbo, Prov. Chapare, Cochabamba, Bolivia, 2,200 meters and
two males from Alto Palmer, Prov. Chapare, Cochabamba, Bolivia, 1,100 meters )
were determined and examined by Richard M. Fox before his death in 1968. He
stated that they were the only known specimens outside of the type series and con-
tained the only known females.

In 1967 I received three additional specimens: a female from Cristal Mayu, Prov.
Chapare, Cochabamba, Bolivia, 600 meters (the lowest known elevation) and two
males from El Limbo, Prov. Chapare, Cochabamba, Bolivia, 2,200 meters, March
1967 (the only known specimens with collecting dates ).

The genus Patricia Fox is better known as Athesis Kirby following usage in Seitz
(1910, Die Gross-Schmetterlinge der Erde, vol. 5). Fox considers Athesis monotypic,
containing only clearista Doubleday & Hewitson; with dercyllidas and allies, includ-
ing demylus, being removed to Patricia. The genus consists of three very rare species
with Andean distribution in Colombia, Peru, Ecuador and (demyllus only) Bolivia.

I have placed a pair of my specimens in the Carnegie Museum collection in
Pittsburgh, and two males in the collection of Herman Real at San Mateo, California.
The remainder, for the present time, are retained in my personal collection.

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin.

FIRST RECORDS OF BOLORIA EUNOMIA (NYMPHALIDAE)
IN WISCONSIN

The first record of Boloria eunomia (Esper) was incorrectly reported by me in
the News of the Lepidopterists’ Society (Number 3, 1969). The correct data for
this record, consisting of two males and one female, is 23 June 1968, in a bog along
highway 111 between Catawba and Phillips, Price County, Wisconsin—instead of,
as originally reported, Rusk County, Wisconsin.

1970 collecting yielded some additional Wisconsin localities for Boloria eunomia.
Fay Karpuleon, of Eau Claire, collected a short series in a bog near Comell, Chippewa
County on 5 June 1970 and added additional specimens during the following week;
this is the southernmost locale where this species has been taken in North America
east of the Rockies. On 14 June, I joined him and we collected additional specimens
(still fresh) in the bog at Cornell, but attempts to locate specimens in bogs further
north were unsuccessful. On 27 June 1970, I secured a few specimens in three bogs
between Toni and Big Falls Flowage in Rusk County and in a bog along highway M
in extreme southeastern Sawyer County, all of which were somewhat flown. Later
the same day, I collected a very wom female in the Comell bog in Chippewa County,
thus documenting a flight season of at least 22 days for the species at this particular
locality in 1970. The flight season for Boloria eunomia is usually regarded as quite
short (e.g. Gray, 1965, J. Lepid. Soc. 19: 184-185).

The range of Boloria eunomia is probably more extensive in Wisconsin than these
scant records indicate. I have found, in Minnesota, that it is one of the more wide-
spread, bog-restricted butterflies, and I expect that it occurs throughout northern
Wisconsin in sphagnum bogs. The Wisconsin population of Boloria eunomia belongs
to the subspecies dawsoni (Barnes & McDunnough).

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin.

150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

ATTRACTION OF SPEYERIA APHRODITE TO ULTRA-VIOLET LIGHT

Hocking Hills is a combinative park, forest and conservation area in southern Ohio,
approximately 60 miles from Columbus. In 1968, 1969 and 1970, from the beginning
of June through August, Lepidoptera were collected by means of long wave ultra-
violet light. The four lights, of the “portable safari-type” were placed five feet
apart and faced in opposite directions. Collecting began at dusk and continued until
approximately 5:00 a.m. on misty evenings with the temperature not below 60 de-
grees F. nor above 85 degrees F. During 1968 and 1969 occasional Papilio polyxenes
asterius (Stoll) and Papilio glaucus (Linnaeus) were attracted to the ultra-violet,
while no specimens of Speyeria aphrodite (Fabricius) were so attracted; all three
species were readily abundant during the day.

In contrast, each collecting night in 1970 yielded four to nine S. aphrodite (sexes
about evenly distributed) at the light, although few S. aphrodite were observed dur-
ing the day. Concurrently, no specimens of P. polyxenes asterius or P. glaucus were
attracted to the ultra-violet light, but these were abundant during the day. The
authors find these observations intriguing. Perhaps further investigation of the at-
traction of diural Lepidoptera to various wave lengths of light would be profitable.

M. S. Cannon AND G. A. Patxuti, Dept. of Anatomy, Ohio State University,
Columbus.

OENEIS JUTTA (SATYRIDAE) IN WISCONSIN

There have been no satisfactory published records for Oeneis jutta Hubner in
Wisconsin. F. R. Arnhold recorded it (Season’s Summary for the Lepidopterists’ So-
ciety for 1954) from Chippewa Falls, Chippewa County, which is further south
than it would be expected to occur; and Masters and Sorensen (1968, Ent. News,
79: 82) referred to specimens from Hines County, Wisconsin; these were specimens
in the Frank Chermock collection, presumably collected by L. Griewisch, but un-
satisfactory as records because there is no Hines County in Wisconsin.

During 1969 and 1970, Fay Karpuleon of Eau Claire, Wisconsin and I collected
examples of Oeneis jutta from a number of localities in Wisconsin including: Chip-
pewa County, near Cornell, 5, 8, 14 and 27 June 1970; Oneida County, near Rhine-
lander, 7 June 1970; Price County, near Catawba, 20 June 1969; and Rusk County,
near Big Falls Flowage 14 and 27 June 1970, near Bruce 20 June 1969, and near
Toni 14 June 1970. In addition, Keith S. Brown Jr. of Rio de Janeiro reports (in
personal communication ) having taken Oeneis jutta in Forest County, Wisconsin dur-
ing 1962.

Oeneis jutta is a bog obligated species in Wisconsin and surrounding areas, and
should have a widespread occurrence in the black spruce/sphagnum bogs of northern
Wisconsin. The species has a biennial life cycle and the heaviest adult flights are
expected in odd-numbered years as in Minnesota. The Wisconsin population of
Oeneis jutta belongs to subspecies ascerta Masters & Sorensen.

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin.

SOME SPHINGIDAE OF HONDURAS

Most of the thirty-four species of Sphingidae listed in this article were collected
by the author from 1968-1970 in Honduras, Central America. All the Sphingidae
were taken from the following five locations: (1) La Ceiba, (2) San Pedro Sula,
(3) La Lima, (4) Tegucigalpa, and (5) Zamorano. The thirty-five species have
been listed alphabetically and numbered according to the locations where they were
collected. See Map of Honduras (Fig. 1) for approximate location of these sites.
Mr. William E. Sieker of Madison, Wisconsin determined the Sphingidae listed.

VoLUME 25, NuMBER 2 1S

a Caribbean Sea
Guatemala >

El Salvador

ae

Nicaragua

Fig. 1. Map of Honduras showing the locations where the Sphingidae were col-
lected. Locations indicated: 1, La Ceiba; 2, San Pedro Sula; 3, La Lima; 4,
Tegucigalpa; 5, Zamorano.

The five locations where the Sphingidae were collected are described below:

La Ceiba. la Ceiba is a seaport of the Caribbean Sea. All my specimens were
taken the nights of July 28, 29, and 31, 1970. All specimens were caught in ‘Colonia
El] Sauce’, being attracted to the big street lights. They were most active between
8:00 and 9:00 p.m. This ‘colonia’ or residential area at the edge of the city, is near
a river and bordered by thick vegetation. The night of July 31 it drizzled around
9:00, and the Sphingidae and large noctuids were very active. After flying around
the lights they would apparently tire and drop to the grass and pavement where
they became “easy picking.”

San Pedro Sula. San Pedro Sula faces the rich Sula Valley with its back to the
mountain. The Sula Valley contains plantations of bananas, sugar cane, and citrus
groves. Most of my Sphingidae were taken at the University which is located outside
the city and surrounded by wooded areas. The sphingids were attracted to the lights
and easily picked off the windows and walls in the evening.

La Lima. La Lima is located 14 kilometers southeast of San Pedro Sula in the
Sula Valley. My specimens were taken at the tennis courts of the United Fruit
Company on August 26 1970 between 8:00 and 9:30 p.m. The bright lights attracted
quite a number of sphingids. A river flows past the tennis courts.

Tegucigalpa. Tegucigalpa is a highland plateau, 3200 feet in elevation, sur-
rounded by pine forest. The Tegucigalpa specimens were attracted by street and
building lights. Many were taken from a high wall along the Choluteca River.
Those too high to reach with a net I knocked down by throwing the sheath of a
hunting knife at them. When touched they would drop straight down into the
awaiting open net.

152 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

Zamorano. The few specimens I have from Zamorano were caught by students of
the Pan American Agricultural School. This school is located in a valley 36 kilometers
east of Tegucigalpa. It has several cloud forest peaks around it. The Sphingidae
were attracted to the lights of the buildings.

The specimens collected are listed below:

Species Location No. Species Location No.
Amphimoea walkeri Boisd. 1 Phlegethontius rustica Fabr. 1, 4
Celerio lineata Fabr. i Phlegethontius sexta Johan. 4
Cocytius duponchel Doey 1 Pholus auchemolus Cramer 4
Epistor ocepete Linné 4 Pholus capronnieri Boisd. 5
Erinnyis alope Drury 1 Pholus labruscae Linné 1
Erinnyis crameri Schaus if Pholus ogliguus R. & J. 1
Erinnyis ello Linné iL, & Pholus vitis Linné IL By 4
Erinnyis lassauxi Boisd. 1 Protambulyx strigilis Linné 2, 4
Erinnyis oenotrus Stoll. iL, B Pseudosphinx tetrio Linné 4
Grammodia caicus Cramer 3 Sphinx merops Boisd. 2,
Hemeroplanes parce Fabr. 4 Xylophanes libya Druce 4
Herse cingulata Fabr. 1 Xylophanes chiron
Pachylia ficus Linné I, WA, 4! nechus Cramer 1
Pachylia resumens Walker 2, al Xylophanes neoptolemus Stoll 1
Phlegethontius florestan Stoll. 1 Xylophanes pluto Fabr. 4
Phlegethontius incisa Walker 4 Xylophanes porcus
Phlegethontius muscosa R. & J. 4 continentalis R. & J. 4
Phlegethontius occulta R. & J. 2 Xylophanes tersa Linné Dy Sw

Rosert D. Leaman, Route 2, Orrville, Ohio.

BOOK REVIEWS

Tue Insect REALM, by Charles L. Hogue and Fred S. Truxal. 1970, 99 pp. + i-viii.
Los Angeles County Museum of Natural History. Price $2.00 U.S.

This attractive little book was nominally produced to serve as a guide to the Hall
of Insects at the Los Angeles County Museum. Such an abundance of information
is presented in concise form, however, that the book could well serve as a text or
reference in high school biology classes. Succesive chapters are devoted to the
position of insects in the animal kingdom, to morphology and phylogeny, to growth
and development and to the insect environment. Other sections deal with beneficial
and injurious insects and the making of an insect collection. The chapter on classifica-
tion gives brief and illustrated diagnoses of the principal insect orders. The book
should be owned by all neophyte entomologists.

D. F. Harpwick, Editor.

A. Fretp Gume TO THE BUTTERFLIES OF BRITAIN AND Europe, by L. G. Higgins and
N. D. Riley, with color illustrations by Brian Hargreaves. Collins, London. 380 pp,
371 maps, 60 colored plates. September, 1970. 42s.

It has been 85 years since there was published in English as complete a study
of the butterflies of western Europe and the British Isles as is presented in this
splendid volume. I am sure that it will be the standard guide to those butterflies

VOLUME 25, NuMBER 2 153

for the rest of this century and well into the next. The book follows the pattern first
established many years ago by Tory Peterson in his bird guides for North America.
The layout is the same as you find in Klots’s Field Guide for the butterflies east of
the Great Plains.

The two authors of the new guide are among the most able Lepidopterists in the
world. Higgins’s careful studies of Old World Melitaeinae brought order to a most
confusing array of butterflies. His personal collection of European butterflies is the
best among all privately held collections. In fact, of the over 720 butterflies illustrated
in this volume 700 are from Higgins’s collection. Dr. Higgins personally collected
by far the major part of his cabinet. Thus he knows intimately the habits and be-
havior of the species within the fauna about which he has written. Riley has been
associated with the butterfly collections of the British Museum (N. H.) since 1911!
He served as Keeper of the Department of Entomology from 1932 to 1955. He edited
the Entomologist for 36 years. No other book about butterflies has behind it the
amount of wisdom that guarantees this one.

The families are arranged in a manner that was in vogue some decades ago and
still is considered the proper ordering by most amateur European collectors. While
it differs from that used in the Zoological Record and from dos Passos’s inversion of
that order, it is no less useful. It is the same as that used by Holland in his Butterfly
Book.

The authors were faced with a fantastic task winnowing the hundreds of sub-
specific names that have been proposed for European butterflies. They settled upon
retaining only those that represent well-defined taxa and eliminated the minor local
varieties. Nowhere is this better shown than in their treatment of the genus Parnas-
sius. They devote two and a half pages to the genus and reduce this far over-named
group of butterflies to seven subspecies of apollo, two of phoebus and two of
mnemosyne.

Careful reading of this book shows that there are striking differences between
the butterfly faunae of western Europe and North America north of Mexico. Part of
this may be related to the more uniform climate of the Old World area when com-
pared with that of the New World area. If the volume considered its eastern bound-
ary the Ural Mountains instead of excluding the U.S.S.R., except for the Baltic states,
more environmental diversity would have been included. The exclusion is reasonable
since few if any western Europeans will be at liberty to collect freely in U.S.S.R. A
summary of faunal differences is presented below:

Group Western Europe U S and Canada
Papilionidae ll 2.9% 28 41%
Pieridae 4] 10.8% 59 8.8%
Danaidae 2 0.5% 5 0.7%
Libytheidae 1 0.3% 2 0.3%
Nymphalidae 68 17.9% 147 21.5%
Satyridae 113 29.8% 47 6.9%
Nemeobiidae Il 0.3% Riodinidae 19 2.8%
Lycaenidae 101 26.6% 133 19.5%
Hesperioidea 4l 10.8% 242, 35.4%

379 99.9% 682 100.0%

Notice that there are many more satyrids in Europe than there are skippers. The
reverse is true in the North American area. Satyrids and skippers compete for
grasses as food in the larval stages. This difference is made more striking when we
compare Hesperiinae, the grass-feeders, in the two areas. We have 128 species, the
Europeans only 12. We cannot blame restriction of grasslands or prevalence of
farmed lands for the small number of Hesperiinae in Europe. The satyrids prove that

154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

there is ample food for a large and varied population of grass-feeders. I suspect that
it is just a case of competition that has existed for many millenia and probably relates
to refugia during the ice advances during the Pleistocene Epoch.

The European subregion is cut off from the tropical faunal regions by deserts and
extraordinarily high mountains. This might be used as an explanation for the some-
what smaller European nymphalid array than is found in America north of Mexico.
Here there is continuous land connection between the tropics and the temperate areas.
The north-south pattern of American mountain ranges also may influence this differ-
ence. At first glance the Lycaenidae seem to deny such reasoning. When this family
is looked at from the subfamily rank quite a different picture developes:

Subfamily Western Europe U. S. and Canada
Theclinae 16 15.8% 80 60.2%
Gerydinae 0 1 0.7%
Lycaeninae Il 10.9% 16 12.0%
Plebejinae 74 13.39% 36 27.1%

101 100.0% 133 100.0%

The dominance of Theclinae in the United States and Canada and the dominance
of Plebejinae in western Europe make the species arrays of the two regions quite
different. Continuity with the tropics, the stronghold of Theclinae, probably explains
the large number of hairstreaks in our fauna. The very large number of blues in the
European fauna may be real or it may be a figment of taxonomic philosophy in the
two listings—Higgins & Riley vs. dos Passos. In turn, each of these is strongly in-
fluenced by recent regional taxonomic research. Stempffer and others in Europe are
far ahead of anyone in North America in understanding of the Plebejinae. I have a
strong suspicion that when parity of intelligence is reached for this subfamily the
number of North America species will be increased. We have tended to turn to
subspecies designation whereas the Europeans have demonstrated specific distinctions
among taxa that are superficially much alike. Perhaps more than a little of our error
has been blind acceptance of work published. We need much more to inquire
critically before accepting the work of others.

Notice of occurrence in North America is included among the brief notes on range
for the holarctic species found in western Europe. Such notice is made for 38 species.
In four cases the relationship is not usually recognized in North America: Pontia
chlorodice beckeri W. H. Edwards, Pontia callidice occidentalis Reakirt, Euchloe
ausonia ausonides Boisduval and Everes argiades comyntas Godart. The last of these
needs verification. Three species in the European fauna which we acknowledge
in ours are not noted as such in Higgins & Riley. These are Lycaeides argyrognomen
Bergstrasser, Vacciniina optilete Knoch and Agriades glandon de Prunner. I wrote
to Higgins about these cases and he replied that in the case of the first two he had
been unable to dissect North American specimens and therefore omitted reference
to them. The omission of glandon was accidental. Higgins had collected material
in Hall Valley, Colorado, when he visited me some years ago. At that time he re-
marked upon how close to glandon is rustica.

Although written primarily for the amateur collector in Europe this book will be
found of value to collectors in North America. I recommend it highly to anyone who
is interested in taxonomic studies, zoogeography and the biology of butterflies.

fF’. Martin Brown, Fountain Valley Rural Station, Colorado Springs, Colorado.

CORRECTION

Vol. 24, no. 4, page 254, line 27: Under Zamagiria australella (Hulst) read “The
type is in the American Mus. Nat. Hist.” for “The type is in the U.S. Nat. Mus.”

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
for material in excess of 20 printed pages, at the rate of $17.50 per page. Address
all correspondence relating to the Journal to: Dr. D. F. Hardwick, K. W. Neatby
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Contributors should prepare manuscripts according to the following instructions;
failure to do so will result in unnecessary delay prior to publication.

Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
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content, including the family name of the subject, but must be kept as short as
possible. The first mention of a plant or animal in the text should include the
full scientific name, with authors of zoological names. Underline only where italics
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Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
_ heading Lirerature Crirep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: All photographs and drawings should be mounted on stiff, white
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Figure legends must be typewritten, double-spaced, on a separate page (not attached
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Proofs: The edited MS and galley proofs will be mailed to the author for
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Tables: Tables should be numbered consecutively in Arabic numerals. Headings
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Material not intended for permanent record, such as current events and _ notices,
should be sent to the editor of the News: Dr. C. V. Covell, Dept. of Biology, Univer-
sity of Louisville, Louisville, Kentucky 40208.

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyn F. pos Passos
Price, postpaid: Society members—$4.50, others—$6.00; uncut,
unbound signatures available for interleaving and private binding,

same prices; hard cover bound, add $1.50. Revised lists of the
Melitaeinae and Lycaenidae will be distributed to purchasers free.

ALLEN PRESS, INC. eRinreo LAWRENCE, KANSAS
usm

CONTENTS

Byers, G. W. A migration of Kricogonia castalia (Pieridae) in northern

Mexico: 2 i I a 2a
Cannon, M. S. and Palkuti, G. A. Attraction of Speyeria aphrodite to

ultra-violet Tight: 0000
Gatrelle, R. R. Notes on the confusion between Lethe creola and Lethe

portlandia (Satyridae) eee

Gatrelle, R. R. Notes on the occurrence of two rare Lepidoptera in South
Carolina 0300 ey a

Gray, R. E. An unusual variant of Colias philodice (Pieridae) in New

Hampshire (ek) |

Hardwick, D. F. The life history of Schinia cupes deserticola (Noctuidae)
Lehman, R. D. Some Sphingidae of Honduras
Manley, T. R. Dragonfly attacks Limenitis defending its territory —__

Masters, J. H. First records of Boloria eunomia (Nymphalidae) in
Wisconsin \ ich

Masters, J. H. Additional records of Patricia demylus gemellus Fox
(Ithomiidae) 2000

Masters, J. H. Oeceneis jutta (Satyridae) in Wisconsin —
Mather, B. The occurrence of Vanessa cardui in Mississippi and. Tennessee

McFarland, N. A specialized case of communal roosting in Pieris rapae
(Pieridae) 2

Munroe, E. A new name for Papilio zetes Westwood, 1847, nec Linnaeus,

1758 (Papilionidae) 00

Nicolay, S. S. A review of the genus Arcas with descriptions of new species
(Lycaenidae, Strymoninit) 200000 a

Oliver, C. G. A melanic aberration of Phyciodes tharos (Nymphalidae)

Opler, P. A. Biological and systematic considerations on the “emarginana
group” of the genus Epinotia (Tortricidae) —...--

Peters, J. V. The Painted Lady Butterfly Vanessa kershawi (Nymphal-
idae), of Australia and New Zealand 00000000) a

Rindge, F. H. Van Someren butterfly collection to the American Museum
of Natural) History) 2050) ON ke

egapaiuertce eisai on Papilio aristodemus ponceanus (Papili-
oniaae

Sevastopulo, D. G. Remarks on “Distributional notes on the genus Mestra
(Nymphalidae) in North America”

Book Reviews

Correction

145-146
143

108
108-114
150-152
146-147

149

149
150
147-148

144-145
142

87-108
143-144

115-123
114
143

126-136

137-139,

139-142

146

154

a

Volume Aa 1971 Number 3

JOURNAL

Leprpoprerists’ SocieTy

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

20 August 1971

THE LEPIDOPTERISTS’ SOCIETY

EDITORIAL COMMITTEE

D. F. Harpwicx, Editor of the Journal
C. V. Covet, Editor of the News
S. A. Hesse., Manager of the Memoirs

EXECUTIVE COUNCIL

C. L. Remincton (New Haven, Conn.) President

Lioyp M. Martin (Prescott, Ariz.) President-elect

H. A. FREEMAN (Garland, Texas) Ist Vice President
Juan Jumaxon (Cebu City, Philippines) Vice President
K. W. Puuie (Fairbanks, Alaska) Vice President

S. S. Nicouay (Virginia Beach, Va.) Treasurer

J. C. Downey (Cedar Falls, Ia.) Secretary

LEE D. Miter (Sarasota, Fla.) Secretary-elect

Members at large (three year term): E. C. Wexiine (Merida, Mexico) 1972
A. E. Brower (Augusta, Me.) 1971 ANDRE BLANCHARD (Houston, Texas) 1973
W. C. McGurrin (Ottawa, Ont.) 1971 R. B. Dominick (McClellanville, S. C.)
Y. NEKRUTENKO (Kiev, U.S.S.R.) 1971 1973
B. Matuer (Clinton, Miss.) 1972 J. P. Donauue (Los Angeles, Calif.)
M. Ocata (Osaka, Japan) 1972 1973

The object of the Lepidopterists’ Society, which was formed in May, 1947 and
formally constituted in December, 1950, is “to promote the science of lepidopterology
in all its branches, . . . to issue a periodical and other publications on Lepidoptera,
to facilitate the exchange of specimens and ideas by both the professional worker
and the amateur in the field; to secure cooperation in all measures” directed towards
these aims.

Membership in the Society is open to all persons interested in the study of
Lepidoptera. All members receive the Journal and the News of the Lepidopterists’
Society. Institutions may subscribe to the Journal but may not become members.
Prospective members should send to the Treasurer full dues for the current year,
together with their full name, address, and special lepidopterological interests.
In alternate years a list of members of the Society is issued, with addresses and
special interests. There are four numbers in each volume of the Journal, scheduled
for February, May, August and November, and eight numbers of the News each year.

Active members—annual dues $10.00
Student members—annual dues $5.00
Sustaining members—annual dues $20.00
Life members—single sum $150.00
Institutional subscriptions—annual $15.00

Send remittances, payable to The Lepidopterists’ Society, and address changes —

to: S. S. Nicolay, 1500 Wakefield Dr., Virginia Beach, Virginia, 23455.

The Lepidopterists’ Society is a non-profit, scientific organization. The office of
publication is Yale University, Peabody Museum, New Haven, Connecticut 06520.
Second class postage paid at Lawrence, Kansas, U.S.A. 66044.

— =

JOURNAL OF

Tue LepiporrTreERISTSs’ SOCIETY

Volume 25 1971 Number 3

The Lepidopterists’ Society, Presidential Address
THE LEPIDOPTERA IN SCIENTIFIC RESEARCH

Pb SHORD Enea S:
Oxford, England

Read by Dr. J. W. Tilden at 21st Annual Meeting of
the Lepidopterists’ Society in Carson City, Nevada
June 21, 1970

There are several reasons why the Lepidoptera provide material es-
pecially well suited to certain aspects of scientific research, in particular,
perhaps, those within the field of ecological genetics. In the first place,
one must mention the beautiful colour patterns of the imagines, which
make it possible to analyse many components of their variation with
especial ease and efficiency. Secondly, and no doubt originating from
their striking aesthetic charm as well as their dramatic metamorphoses,
butterflies have been one of the two animal groups most studied by gen-
erations of naturalists (birds being the other). Thus it comes about that
we possess an immense fund of information on their ecology; these insects
really have attracted more attention and for a longer period than any
others, and the knowledge so obtained is of practical importance as a
background for many types of biological research. It has indeed grad-
ually accumulated until Lepidopterists have found the need to establish
an International Society such as ours, in which the old natural history is
transformed into an important branch of modern science. Yet let us not
forget the entomologists of an earlier age. Some who were collecting
butterflies and moths 120 years ago have been my personal friends and I
know what fine field workers they were: their store of information,
largely centred upon habit and behaviour, for the most part died with
them, but it was great.

When we consider the impact of the Lepidoptera upon scientific re-
search, one point strikes us at the outset or else is accepted as a basis
inherent in the material we use: that is to say, their taxonomy. For if

156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

we are to study any large assemblage, of whatever kind, it is necessary to
classify it hierarchically. In doing so we must create some system giving
us confidence in the names applied to the groups we use. It is perhaps
invidious to single out any of those who have laboured to establish the
nomenclature of the Lepidoptera. However, I would at least mention Mr.
Francis Hemming, whose death some years ago was so serious a blow to
this work, and Dr. C. F. dos Passos, of our own Society, to whose studies
in this field at the present time taxonomy owes so much.

But what of the groups which the taxonomist names? We are all aware
that though many possible classifications could be applied to them, that
in general use is based upon relationship, so that it reflects one im-
portant aspect of reality; that is to say, evolution. It is our aim that the
Lepidoptera placed in any one taxonomic division, whether it be small
or great, species or family, should be more closely related to one another
by actual descent than they are to any other organisms upon earth. Thus
if the arrangement be represented diagrammatically, the result should pro-
duce a geneological tree.

We should certainly enquire whether that object, so basic to our en-
deavours, has been attained. Is the classification of butterflies and moths
something more than a man-made convenience; does it indeed reflect
affinity? We have a fair guarantee in that matter arising from the work
of taxonomists themselves, who generally take into account the widest
range of characters in making their decisions: morphological and physio-
logical and in all the stages of the life history. But we can test its success
when we extend to it a new criterion. We shall be on sure ground if in-
dependent evidence supports conclusions reached in ignorance of it.

Many years ago it seemed to me worthwhile to seek such confirmation.
I obtained it by examining the chemistry of the wing pigments of butter-
flies and moths. These indeed proved to be strictly related to a classi-
fication which had taken no account of them. We have time to cite one
instance only. Consider the flavones: these are non-nitrogenous pigments,
ranging from deep yellow to ivory-white and derived ultimately from
plants. Certain Lepidoptera make use of them to colour their scales,
others do not. But those which employ them in that way are not scattered
at random among the rest. Thus in the family Pieridae, flavones are
never found except in the sub-family Dismorphiinae, and even there in
but a proportion of the species (19 out of 58 tested: from a total nom-
inally assessed as 108). Yet, with one exception independently admitted
to be a taxonomic error, those in which they occur had been grouped
together, although this particular quality which they share was unknown

VoLUME 25, NUMBER 3 WL5S7/

to the entomologists who classified them (Ford, 1941); many other similar
corroborations have subsequently been obtained.

Bearing these advantages in mind, the Lepidoptera have naturally been
extensively used in scientific research, and a few examples of the prob-
lems they have helped to illuminate may briefly be considered here. We
may first take the question of selection-pressure, and indicate two of the
instances in which it has been evaluated.

The butterfly Maniola jurtina L., which is single brooded in Britain
and most of Europe, has provided information on this subject. A criterion
of that insect’s adaptive adjustment is supplied by the spots on the under-
side of the hind wings. These may be absent or present in any number
up to five, arranged as a curving submarginal row. They are controlled by
polygenes which influence also other characteristics of greater importance
to the organism and of these the spots may be taken as an outward and
visible sign. Thus they affect the date of emergence, the more spotted
specimens appear earlier, and also affect liability to destruction by certain
parasites. These polygenes moreover, play an important part in adjust-
ing the gene-complex to the needs of the population. When the butter-
flies are reared in a cool laboratory, with a temperature fluctuating in the
region of 15° C, the heritability of spotting is 0.83 in the females and 0.14
in the males (McWhirter, 1969). At higher temperatures, the environ-
mental component, so much greater in the males than in the females, is re-
duced; for at 22° C heritability actually approaches 1.0 in the females
and it reaches 0.4 in the males. Thus selection has much genetic vari-
ability on which to operate.

We may concentrate here upon the female spot-frequencies since,
being subject to greater major variability, they are more diagnostic than
those of the males, which are nearly always unimodal at two spots. Not
so the females, in which the spot-frequencies are diverse and character-
istic.

The Isles of Scilly are an archipelago in the Atlantic 30 miles from the
south-western extremity of England. We have studied Maniola jurtina
upon a number of the Islands there, some large (682 acres or more),
others small (40 acres or less). The difference in area between the two
types is, therefore, at least 17 times. Female spotting differs greatly from
one to another of the small Islands (it may be unimodal at 0 or at 2, or
bimodal with the greater mode either at 0 or at 2), retaining its character-
istic value on each, year after year. It is, however, similar on the major
areas of all the large islands (with approximately equal values at 0, 1 and
2 spots), a frequency not found on the small ones. What, may be enquired,
is the meaning of such distinctions. In interpreting them we have to

158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

remember that the spotting is stabilised at characteristic values on each
island. This in itself excludes an explanation based on random genetic
drift, so does the size of the various populations involved, for they may
comprise thousands of imagines, ascertained by the technique of marking,
release and recapture. We have indeed strong reasons for thinking that
the butterflies are adjusted to the distinct ecology of each small Island
but to the average of the conditions on the much more diversified large
ones, and such averages tend to resemble one another (Ford, 1965).

It has also been suggested that the Founder principle operates here.
Yet changes in these highly constant spotting types have actually been
witnessed as the result of rare cataclysms. They have produced new and
stabilised spot-frequencies differing as much from the old as those on
one small Island differ from another.

We may consider the situation on White Island, which is divided into
two parts, each about a quarter mile long, by a low narrow neck of
land. This was covered by sand and shingle during a great storm in the
winter of 1957-58. As a result, the Maniola jurtina population was sub-
divided. That on the southern half of the Island, previously stabilised
with a mode at no spots, changed at once to a condition unimodal at one
spot and remained for eleven years at the new frequency; until in 1969
the recently created barrier ceased to be effective owing to the growth
of plants upon it and the spotting returned to its pre-1958 values. A
selection-pressure of 68 per cent with 5 per cent confidence limits at 31-
81 per cent, in favour of one-spotted individuals was needed to produce
that result in 1958 (Creed e¢ al., 1964).

Turning briefly to Maniola jurtina as found in south-west England,
the females there are bimodal at 0 and 2 spots, though in the rest of the
country and throughout Continental Europe, except in its more peripheral
regions, they are unimodal at 0 (Dowdeswell & McWhirter, 1967).
Though the boundary between the two frequencies has repeatedly altered
its position by several miles in a single generation of the butterfly, one
type changes into the other in a few yards; it does so along a line crossing
the peninsula of Devon from north to south. No physical boundary is in-
volved and imagines of the respective stabilisations are constantly inter-
mingling where they meet (Creed et al., 1959).

In 1957 the boundary between the bimodal and unimodal types moved
three miles eastward. This involved selection of 65 per cent, with 5 per
cent, confidence limits of 31 to 82 per cent, against two-spotted in-
dividuals in the population involved. Such high selection-pressures now
prove to be usual in the micro-evolution of the Lepidoptera, as we find
them to be in other organisms.

VOLUME 25, NUMBER 3 159

In addition to selection, one of the features operating in evolution is
provided by isolation, as Darwin clearly points out. An aspect of this, a
curious one seldom considered, has just been mentioned: that provided
by selection so powerful that it can break up a population into two ad-
justed types, maintained without geographical discontinuities, a situation
sometimes manifested by the formation of a reverse cline (Ford, 1965),
as indeed in Maniola jurtina. That is to say, the distinguishing character-
istics of two populations become accentuated towards the line where they
meet; selection being powerful enough to eliminate the less well adjusted
intermediate types at the interface between them. Since, as we have
seen, one spot-adjustment can be converted into another, there is no
question of a past gap in the distribution of this insect from which pop-
ulations with distinct spotting-types have spread until they have come
into contact. That, of course, can occur and it produces somewhat similar
results to the one just mentioned, though not the conversion of a part of
one race into the other. Thus the Scandinavian and European races of
the Selidosemid moth Bupalus piniarius L. meet and interbreed across
England at the level of Lancashire and Cheshire (Cockayne, 1912-13).
The one must be an Ice Age relict which has retreated northwards, the
other a Holocene colonist from Europe. Yet their approximation has not
produced a cline. On the contrary, there is an area of high variability
between these differing adjacent populations, within which the re-
combinants are constantly produced though they fail to spread in either
direction so as to swamp the characteristics and balanced adaptations of
the two forms.

We have not indeed extensive information on the genetics of speciation
and racial differences in the Lepidoptera. The distinctions between the
Bupalus piniarius races appear to be multifactorial. Those separating
closely related species may be also, or they may include characters re-
sulting from the action of major genes.

The Lycaenid genus Aricia contains two European species not separated
until 1935. These are the bivoltine A. agestis Schiff., the northern limit of
which is reached in southem England and northern Denmark, and the
univoltine A. artaxerxes F. from northern Britain, Norway, Sweden and
Finland (Frydenberg and Hgegh-Guldberg, 1966). Each includes a
number of more or less distinct races and their characteristics must not
be confused with those of the species as a whole. For instance, the
features distinguishing artaxerxes artaxerxes from artaxerxes salmacis con-
sist of a white central spot on the upper side of the forewings and an ab-
sence of the central black dot within the white spots on the lower sur-
face. That condition is due to a single gene recessive in effect.

160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

However, a number of characters, none in itself entirely distinctive,
separate A. agestis and A. artaxerxes. These are quantitative only, yet
some are unifactorial: thus one is provided by the more heavily marked
larva of the latter species, another by the larger submarginal orange
lunules on the upper side of the imagines (Hgegh-Guldberg, 1968;
H¢gegh-Guldberg and Jarvis, 1969). No feature is said to be fully diag-
nostic of these two butterflies except the double and single brooded habit.
This is by no means a specific trait in other Lepidoptera; it is not so, for
instance, in the moth Zygaena lonicerae v. Schev. (Lane and Rothschild,
1961). In Aricia that quality seems to depend on combined environmental
and genetic influences, the latter multifactorial. Here, however, it does
seem to some extent to separate two groups specifically, for crosses be-
tween them are to some extent infertile.

A rather similar situation from the evolutionary point of view is pro-
vided by Pieris napi L. and P. bryoniae Ochs. It is now said that their
chromosome numbers differ: n = 25 in napi and 26 in bryoniae ( Bowden,
19662). P. napi is Holartic and occurs from sea-level up to high altitudes.
P. bryoniae is alpine and boreal only, having a discontinuous distribution
which includes parts of Scandinavia, the Alps, Central Asia and Canada.
The two insects are separated by a number of features which tend to
maintain their reproductive isolation: habitat, differences in time of
flight, low viability of the hybrids and differential mating behaviour
(Petersen, 1963). Their larval foods are not the same and though the
males are very similar the females are highly distinct. Bryoniae differs
from napi principally owing to the action of several major genes (Lork-
ovic, 1962): (1) B, which is autosomal and nearly dominant in effect
spreading melanin along the veins, and (2) the gene Y which produces
the ochreous-yellow ground-colour of bryoniae. It is incompletely domi-
nant and is almost entirely sex-controlled to the females.

In addition, many populations of bryoniae, especially in the eastern
Alps, are polymorphic for a gene W producing dominant white coloration
on the underside of the hind wings and tips of the forewings (Bowden,
1963). This is allelic with that responsible for the two recessive yellow
phases (sulphurea) found as rare recessive mutants in P. napi. It has
been said that the polymorphism of this white-underside form (subtalba)
is partly maintained by the lethality of the homozygotes. Bowden (1967)
produced some evidence for this but certainly obtained homozygous
whites. In this matter, a point of general importance must be recognised.
That is to say, a selective disadvantage of 10 or 15 per cent may be effec-
tive in maintaining a polymorphism but very difficult to detect in experi-
mental breeding.

VOLUME 25, NUMBER 3 161

It is especially to be noticed that both the genes Y and W, and probably
B, occur as rare mutants in the normal population of P. napi (Bowden,
1963). We have here a clue to the extremely discontinuous distribution of
bryoniae. That is to say, the genes capable of producing it are widely
available when their action is appropriate to the ecology of this Pieris
complex.

There are, of course, many instances in which a form controlled on a
unifactorial basis appears to recur in isolation, though we do not know
whether the same gene be responsible for it in the different habitats.
Thus the Arctiid Cycnia mendica Cl. is ordinarily a sexually dimorphic
species with black males and white females bearing black dots. In the
Irish race, rustica, the males are white like the females, due to the action
of a gene sex-controlled in effect and giving intermediate heterozygotes
(Onslow, 1912; Ford, 1967a, plate 10). Yet rustica recurs in a few pop-
ulations in Europe, or else its phenotype does so.

The moth Triphaena comes Hb., Caradrinidae, provides one of the few
instances in which a phenomenon of this kind has been analysed from a
comparative point of view. The dark form, curtisii, occurs as a polymor-
phism in three isolated populations: the Outer Hebrides, the Highlands of
Scotland and in Orkney. It is identical in appearance in all of them. It has
been shown that the Hebridean and Orcadean populations are due to the
action of one major gene and that it is the same in both places, but it is
adjusted by different modifiers to give an apparently identical effect in
each (Ford, 1955). Here we really do know what is meant when we say
that the “same” form of the species occurs in two distinct areas.

From this point we are, by an easy transition, taken over to another
system of variation, that of polymorphism, and to another aspect of evolu-
tion, that of Industrial Melanism.

The Lepidoptera are, of course, famous for providing that outstanding
reaction to pollution. Though known in other groups, (e.g. Adalia bi-
punctata L. among the Coleoptera, Coccinellidae), it is in the moths that
Industrial Melanism is most widespread and striking (Kettlewell, 1957,
1961). This is not the place in which to discuss that situation, for it
merits long and detailed attention, but a few of the facts relating to it can
briefly be summarised.

Many dozens of species have become black in the industrial areas of
Britain (where approximately 100 are affected), Continental Europe, the
U.S.A. and elsewhere. Nearly all are controlled by a single major gene,
dominant in effect. They spread owing to the action of two agencies.
One is bird predation of the less cryptically coloured forms: the melanic
insects on light coloured bark and lichens; and the normal pale ones on

162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

trees, fences and other sites which are blackened by pollution in manu-
facturing districts. The second agency responsible for the spread of in-
dustrial melanics is the evolution of heterozygous advantage, which is
brought about in either of two ways or by a combination of them: 1) the
accumulation of common lethal or semi-lethal mutants close to the locus
controlling the polymorphism; 2) the evolution of dominance, operating
in different directions upon the multiple effects of the switch-gene. This
causes the disadvantageous qualities it determines to become recessive
and the advantageous ones dominant, so ensuring the superiority of the
heterozygotes over the other two genotypes.

The occurrence of Industrial Melanism provides an exceptionally sensi-
tive test for the spread of pollution outwards, and down the prevailing
wind, from industrial areas. Moreover, we have here excellent examples
of evolution in progress, for the melanic forms of many moths even
though unifactorially controlled have become progressively better ad-
justed to their environments, both in their physiology and in their colour-
ing, by selection operating on their gene-complexes (Kettlewell, 1965).

Only very rarely is an industrial melanic recessive. The large Bombycid
Lasiocampa quercus L. provides an instance of the kind on a moor in
Yorkshire. The area is heavily polluted with soot and a blackish-brown
form of the moth occurs there (Kettlewell, 1959). Probably it is prefer-
entially selected by the large numbers of Gulls which prey upon the
species in that locality. The blackish specimens at present amount to
only about 5 per cent of the population; though of course the gene re-
sponsible for them must occupy approximately 35 per cent of available
loci, more if the heterozygotes are at an advantage.

An identical melanic is also found occupying up to 70 per cent of the
population on heather moors in northeastern Scotland. There the plants
are stunted and widely spaced, exposing the black peaty soil and giving
the whole area a dark appearance. Here we return to the aspect of isola-
tion that has just been discussed, for this is another of the rare instances in
which it has been studied experimentally. The result proved that the two
apparently similar blackish forms, in Yorkshire and north-east Scotland
respectively, are generically distinct, being produced by different major
genes. That in the Scottish population, moreover, is not fully recessive in
effect.

In the Yorkshire population, where recessiveness is complete, an addi-
tional feature of interest is to be noticed. That is to say, a recessive black
form of the larva also exists, which is absent from the Scotch locality. It
is due to a separate gene from that giving rise to the dark imagines, but

VOLUME 25, NUMBER 3 163

the two loci are closely linked: in fact they are probably being built into
a super-gene.

This indeed is characteristic of co-adapted genes when polymorphic.
Selection tends to produce close linkage between them so that the ap-
propriate features can segregate together. For polymorphism involves dis-
continuous variation, as in the human blood groups, in which even the
rarer form is maintained selectively. The phases are indeed balanced
either ecologically, as to some extent in butterfly mimicry, or by a supe-
riority of the heterozygotes (Ford, 1965). Thus the evolution of super-
genes, as of heterozygous advantage, will generally take place in poly-
morphism which must in view of these attributes be a very common
phenomenon. It is, moreover, one which must nearly always be con-
trolled genetically, by the segregation of a major gene or a super-gene.
Environmental differences are indeed able to affect profoundly entire
broods and populations, as in seasonal variation (which we see, for in-
stance, in the Nymphaline butterfly Araschnia levana L.) (Ford, 1967b).
They are hardly capable, however, of evoking discontinuous variability
throughout a considerable region. This indeed becomes an impossibility
when, as in polymorphic situations, the frequencies of the forms must be
appropriately adjusted to differing ecological situations.

Polymorphism is usual in the Batesian mimicry of butterflies, in which
its properties are clearly seen. For as the relative numbers of mimics
sheltering under their resemblance to a distasteful form rises, so the bene-
fit they obtain from doing so declines until it is lost and converted into a
disadvantage. Therefore selection will favour the tendency for a Batesian
mimic to copy several models. This it must do by means of polymorphism:
evoking a mechanism, that is to say, which avoids intermediates and
and therefore the production of unprotected forms.

Yet the switch-gene necessary to achieve this must have arisen by muta-
tion, and we cannot expect its original effects to have been accurately
adjusted so as to resemble a particular model. That is achieved gradually
by selection, acting upon the gene-complex, within the ambit of the con-
trolling major gene. Thus it is repeatedly found that mimicry even when
“unifactorial” becomes imperfect in crosses with a race in which the ap-
propriate model is absent (Clarke and Sheppard, 1960).

The two major consequences of polymorphism are also exemplified in
mimicry. The phases of a Batesian mimic often include non-mimetic
forms which, as in normal polymorphic situations (the white and yellow
females of Colias, for instance: Remington, 1954), are maintained by
heterozygous advantage. This, and the selective adjustment involved, is
demonstrated by the fact that mimics can sometimes exist even at high

164 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

frequencies outside the range of their models (as in the Polytrophus
race of Papilio dardanus Brown, in the mountains east of Lake Victoria).
That they do so demonstrates their physiological advantage; it demon-
strates also the importance of their deceptive colour-patterns, for the
accurate resemblance of these is lost in such circumstances (Clarke and
Sheppard, 1960b ).

The adjustments of mimicry are usually multiple, requiring the com-
bined action of co-adapted genes. These must be held together in ap-
propriate combinations by close linkage; their inappropriate ones, also
produced, are eliminated by selection in the same way as are ordinary dis-
advantageous mutants. Clarke and Sheppard (1960c) have illustrated
very clearly the evolution of such super-genes in mimicry. In this con-
nection, their work on the Asiatic Papilio memnon L.. is impressive (Clarke
et al., 1968). This species has numerous polymorphic female forms,
some non-mimetic and some mimetic. Of the latter, one (achates), copies
a tailed model (Atrophaneura coon F.), though the others do not, and
it is accordingly provided with a tail. The gene responsible for this is
included within the super-gene for colour-pattern so that it may operate
in the necessary genetic setting, but not otherwise. However, the race
of P. memnon inhabiting the Island of Palawan is exceptional, being in-
variably tailed, a condition also due to the action of a single gene. But
this is not included within the super-gene controlling colour-pattern, for
in this instance wing-shape is unassociated with mimicry.

An interesting discovery has lately been made which, in a sense, bridges
the gap between the Batesian and Miillerian situations. In the latter, a
number of inedible and warningly coloured butterflies, or other insects,
shelter under a common colour-pattern, so that the lesson of distasteful-
ness learned by a predator attacking one of them may apply to all.
Brower, Brower & Corvino (1967), working on Danaus plexippus L., find
that in this instance the objectionable substances present in the imago
are derived from the larval food. When, however, as occasionally hap-
pens, such larvae eat non-poisonous plants, the resulting butterflies lack
their normal protection and so shelter under the reputation of inedibility
gained by the majority of the species, a situation which Brower e¢ al.
name automimicry. It may well involve many species in addition to D.
plexippus.

Variation may be cryptic and, since as a gene spreads it will tend to
generate heterozygous advantage and therefore become polymorphic,
cryptic polymorphism must be widespread and important. Its existence
is now being detected by means of electrophoresis, by which at least a

VoLUME 25, NUMBER 3 165

proportion of the protein-variability present in organisms can be recog-
nised. Appropriate staining techniques can sometimes demonstrate all
three genotypes controlling alternative esterases, and it has repeatedly
been found that the heterozygotes are in excess of expectation, so estab-
lishing their superiority. Lewontin and Hubby (1966) who were pioneers
in this work have indeed attempted to show that such protein diversity
cannot be due to heterozygous advantage. They were lead to that view
by the intrusion into their argument of two errors, well exposed by Milk-
man (1967). First, that the genes controlling distinct polymorphisms act
independently. Yet as Milkman remarks, “artificial selection has taught
us nothing if not the cumulative effect of genes at many loci.” Secondly,
Lewontin and Hubby hold that the unit of selection is the gene, whereas
it is the individual. On the latter, and correct, basis Milkman develops a
simple calculation which demonstrates the power of heterozygous advan-
tage to maintain protein polymorphism.

Johnson and Burns (1966) using starch-gel electrophoresis found that
the butterfly Colias eurytheme Bd. is subject to great protein (esterase )
polymorphism, which differed both quantitatively and qualitatively in
two populations from localities in Texas approximately 100 miles apart.
Indeed, this technique has opened up a number of new possibilities in
the analysis of adaptation and micro-evolution in general.

When in the 1920’s I was envisaging the experimental study of evolu-
tion in wild populations, it seemed necessary to concentrate upon situa-
tions which promote rapid adaptation. Three of these proved highly
satisfactory for that purpose but a fourth, the spread of species into new
territories, was disappointing. I deduced that in the instances which
happened to be available for study at the time, the adjustments which
must surely be taking place were largely physiological. The technique
of electrophoresis now provides an opportunity for testing that view.
Numerous instances of species colonising new lands spring to mind. One
may mention the Palaearctic Pieris rapae L., which has become one of
the commonest butterflies in the U.S.A. since it was introduced there in
the middle of the last century. Has it adjusted its protein variation to live
in the New World? Electrophoresis will probably provide an answer to
that question.

A somewhat exceptional type of polymorphism occurs in a single popu-
lation of the moth Panaxia dominula L., Hypsidae (Ford & Sheppard, 1969).
Its study has produced results which call for some comment. The species
has one generation in the year and is widespread in Continental Europe
and southern England; occurring in marshes and along river banks, where

166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the larvae feed chiefly upon Symphytum. The polymorphism occurs nat-
urally in a single locality only, one of 17 acres in Berkshire. This has
certain peculiar features of soil and ecology which may account for the
exceptional characteristics of the moth there.

Its polymorphism is controlled by a pair of alleles in which all three
genotypes are distinct. It is therefore possible to calculate gene-frequency
by inspection, which has so far been done yearly from 1939 to 1970.
Moreover, the numbers in the colony have been assessed by the technique
of marking, release and recapture from 1941 onwards. It is possible to
determine how fast a gene can spread if both its frequency and the size
of the population in which it occurs be known. In this instance, for the
first time, both these requirements were met. The annual fluctuation in
the ratio of the alleles proved to be too great to be the result of random
processes, such as genetic drift, and must be due to selection changing
sometimes in frequency and direction.

In this instance, the polymorphism is not controlled by heterozygous
advantage; indeed there is a clear indication that there has not yet been
time for this to evolve in the community. The diversity is, in fact, main-
tained by assortative mating, for the females favour pairing with males
of either of the two phases to which they do not belong.

The heterozygotes are somewhat variable. Indeed it proved possible
by only four generations of selection in the laboratory on the one hand
significantly to increase and on the other to diminish their expression, in
the sense of departure from the normal homozygous type: that is to say,
changing the terminology, to make them respectively more dominant or
more recessive.

In 1954 the polymorphism was started artificially by P. M. Sheppard
in another isolated colony a mile away. It is now found that changes
similar in type to those produced in the laboratory, but taking place much
more slowly, are occurring in these two wild communities. The expression
of the gene is becoming more dominant in the marsh where it occurred
naturally and more recessive in the one where it was introduced. This
is perhaps the first time that it has been possible to forestall experiment-
ally an evolutionary change that has occurred subsequently in nature.

The most importance outcome of using the Lepidoptera in scientific
research, as indeed in applying the techniques of ecological genetics to
organisms in general, is surely the discovery that natural selection is
much more powerful than had previously been realised. We had been
accustomed to think of a selection-pressure of 1 per cent in favour of
advantageous qualities in nature as high. Today we realise that it fre-
quently reaches 40 or 60 per cent. Here we must notice that these values

VOLUME 25, NUMBER 3 167

are often balanced in equilibrium, so that the overall selective advantage
or disadvantage may indeed be small. Yet the powerful components in-
volved facilitate rapid adjustments to changing conditions, a situation
which alters fundamentally our concepts of micro-evolution.

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Forp, E. B. 1967b 3rd edn. Butterflies. The New Naturalist Series, Collins, Lon-
don.

Forp, E. B. anp P. M. SHeppArp. 1969. The medionigra polymorphism of Panaxia
dominula. Heredity 24: 561-9.

FRYDENBERG, O. AND O. H6rcH-GuLpBerc. 1966. The genetic difference between
southern English Aricia agestis and Scottish A. artaxerxes. Hereditas 56: 145-
58.

H6rcH-GuLpBErc, O. 1968. Evolutionary trends in the genus Aricia (Lep.).
Natura Jutl. 14: 3-76.

H6EcH-GULDBERG, O. AND F. V. L. Jarvis. 1969. Central and North European
Ariciae (Lep.). Natura Jutl. 15: 1-119.

Jounson, F. M. anv J. M. Burns. 1966. Electrophoretic variation in esterases of
Colias eurytheme (Pieridae). J. Lepid. Soc. 20: 207-11.

168 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

KETTLEWELL, H. B. D. 1957. The contribution of industrial melanism in the
Lepidoptera to our knowledge of evolution. Advmt. Sci., Lond. 52: 245-52.
KETTLEWELL, H. B. D. 1959. New aspects of the genetic control of industrial

melanism in the Lepidoptera. Nature 183: 918-21.

KETTLEWELL, H. B. D. 1961. The phenomenon of industrial melanism in the
Lepidoptera. A. Rey. Ent. 6: 245-62.

KETTLEWELL, H. B. D. 1965. Insect survival and selection for pattern. Science
148: 1290-6.

Lange, C. AND THE Hon. M. Roruscuitp. 1961. Observations on Colonies of the
Narrow-bordered Five-spot Burnet (Zygaena lonicerae von Schev.) near Bicester.
Entomologist 94: 79-81.

LEWONTIN, R. C. anv J. L. Hussy. 1966. The amount of variation and degree of
heterozygosity in natural populations of Drosophila pseudoobscura. Genetics
54: 595-609.

Lorkovic, Z. 1962. The genetics and reproductive isolating mechanisms of the
Pieris napi-bryoniae Group. J. Lepid. Soc. 16: 5-19, 105-127.

McWuirter, K. G. 1969. Heritability of spot-number in Scillonian strains of the
Meadow Brown Butterfly (Maniola jurtina). Heredity 24: 314-18.

MiLkMAN, R. D. 1967. MHeterosis as a major cause of heterozygosity in nature.
Genetics 55: 493-5.

OnsLow, THE Hon. H. 1921. Inheritance of wing-colour in Lepidoptera. J. Genet.
11: 277-92.

PETERSEN, B. 1963. Breakdown of differentiation between Pieris napi and Pieris
bryoniae and its causes. Zool. bidr. Upps. 35: 205-62.

REMINGTON, C. L. 1954. The genetics of Colias (Lepidoptera). Adv. Genet. 6:
403-50.

FERGUSON COLLECTION TO SMITHSONIAN INSTITUTION

Dr. Douglas C. Ferguson, formerly of the Nova Scotia Museum and Yale Univer-
sity, recently assumed a position as specialist on Geometroidea and Pyraloidea with
the United States Department of Agriculture at the National Museum of Natural
History. Shortly after joining the staff, Dr. Ferguson presented to the Smithsonian
Institution his excellent collection of Lepidoptera. Over 51,000 insects are contained
in this collection, of which nearly 48,000 specimens are Lepidoptera. Dr. Ferguson’s
donation represents over 23 years of concentrated collecting, conducted for the most
part in the Atlantic Provinces of Canada. Nearly all major families of Lepidoptera
are represented, and the total coverage of 3,000 species from an area previously not
well documented in the collections of the National Museum is particularly noteworthy.
The collection is extremely rich in Noctuoidea and Geometroidea, containing ap-
proximately 38,000 beautifully prepared specimens of those groups. Over 150 species
of identified lepidopterous larvae, associated with reared adults, are also included.
Although no holotypes are present in the collection, about 300 paratypes of 50 species
are, as well as many undescribed species.

The addition of this generous gift has significantly improved the overall representa-
tion of North American Lepidoptera at the Smithsonian Institution.

DonaLp Ray Davis, Curator, Department of Entomology, Smithsonian Institution,
Washington, D.C.

VOLUME 25, NUMBER 3 169

NEW SPECIES OF AFRICAN GEOMETRINAE (GEOMETRIDAE)

R. H. Carcasson

Centennial Museum, Vancouver, B.C.

Victoria watsonae Carcasson, new species
Figs. 1, 14

Allied to V. melanochlora Carcasson (1962: 54) and to V. gordoni Prout
(1912: 86), but differs from both in having the edges of the black mark-
ings diffuse, not well defined and in having no densely scaled green areas.

MALE. Antennae: Basal half pectinated, distal half simple. Dorsal aspect of shaft
covered by pale pinkish brown (5YR 8/4 Munsell) white and sepia scales. Head:
Vertex pinkish brown speckled with sepia. Frons whitish. Palpi cinnamon with sepia
scales, laterally fringed with white below. Thorax: Patagia and tegulae light blue
green (10G 7/4 Munsell). Dorsum anteriorly and laterally light blue green, reddish
speckled with sepia at base. Venter whitish except for a vinaceous light red anterior
collar. Inner surface of legs whitish, outer surface of tibiae and tarsi cinnamon speckled
with sepia. Abdomen: Above pinkish white, 3 anterior segments heavily speckled
with sepia and cinnamon. Segments 2 and 3 with a prominent dorsal crest consisting
of very long, curly, pinkish cinnamon and sepia scales. Similar crests on 4, 5 and 6,
but much paler, laterally edged with sepia. The crest on 6 just a small white tuft.
Underside creamy white. Upperside of forewing: 17 mm from base to apex. Apex
rather acute, termen regularly indented between veins. Wing lightly scaled, ground
colour light blue green (10G 7/4 Munsell) with a few scattered blackish scales.
Costa brownish pink speckled with sepia. Stigma a ring of raised blackish scales.
Antemedial irregular, reduced to a blackish spot near costa, a faint blackish spot at
cubitus and another at inner margin, connected by irregularly scattered blackish scales.
Postmedial roughly parallel to outer margin, consisting of small irregular blackish dots
at the veins from costa at 4/5 from base to inner margin at 2/3 from base. A large
blackish blotch with diffuse margins containing scattered pinkish scales between post-
medial and termen, from vein 6 to tornus and inner margin, reaching termen except
between veins 3 and 4 where it encloses a large green marginal spot. A black terminal
dot in space 3 and smaller one in space 6. Fringe blackish where dark areas reach the
margin, otherwise pinkish mixed with blackish and green scales. Hind wing: Margin
with deep inter-nervular indentations. Ends of veins 2, 3, 4 and 6 especially produced.
Ground colour as in fw. A small patch of dense, silky white scales at base. Stigma a
densely scaled black dot surmounted by a smaller orange pink dot. Postmedial ir-
regular, consisting of small ill-defined blackish dots at the veins. Inner margin blackish
with a few pink scales, except at base which is green. Tornus and termen up to vein
3 blackish. A blackish dot in space 2 near termen. A small black terminal spot in
space 3, a larger one in 4, some black scales at termen of space 6 and at apex. Fringe
as in fw. but with more green in pale areas. Underside: Thinly scaled, opalescent
white, with dark stigmas, marginal spots and postmedial faintly showing through
from upper surface. Blackish fw blotch as above, but denser, lacking pink scales.

Genitalia: Uncus sharply pointed, only slightly downcurved forming a rough equi-
lateral triangle; a small rounded heavily sclerotised plate, almost at centre of uncus.
Subunci membranous, hairy, not reaching tip of uncus. Gnathos a stout pointed
process directed ventrad attached to a strong, heart-shaped sclerotised ring. Saccus
short and pointed. Valve long and narrow, spatulate, terminating in a dorso-apical
lobe densely covered internally by long, stout peg-like projections. Harpe long, narrow,
sharply pointed, protruding beyond apex of valve. Aedeagus short and stout, lightly
sclerotised, with long longitudinal apical opening on left side. Vesica protruding from

170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-9. African Geometrinae. 1, Victoria watsonae sp. nov., 6; 2, Archichlora
rectilineata sp. nov., 6, holotype; 3, same, 8, paratype (dark form, Mufindi); 4, same
6 (Kigezi, Uganda); 5, same 9, allotype; 6, Archichlora sangoana sp. nov., 6, para-
type; 7, Archichlora jacksoni sp. nov., ¢, holotype; 8, Archichlora pulveriplaga
(Warren), 6 (Kalinzu forest, Ankole, Uganda); 9, Archichlora rectilineata sp. nov.,
2? (Njoro, Kenya. )

All specimens reproduced at natural size.

VoLUME 25, NUMBER 3 eval

left side of aedeagus, armed with two very large curved blunt spines, the apical
larger and directed to the right, the smaller arising well below the apical and curving
towards the left when not extruded.

Holotype. ¢, Grassfield, Nimba, Liberia, VI-VII 1967, Mrs. A. Forbes-Watson,
to be deposited in the British Museum (Natural History). Paratypes: 44 6, data
as above; 1, VI-VII 1967, 1 VII-VIII 1967, 1 II 1968, 1 VIII-IX 1967, to be deposited
in National Museum, Nairobi, Kenya.

Archichlora jacksoni Carcasson, new species
Figs. 7, 10
Allied to A. pulveriplaga (Warren), (1898: 14), (figs. 8, 12), but differs

in its smaller size, continuous pinkish postmedial line and in the presence
of a basal pinkish blotch in the forewing.

MALE. Antennae: Basal half armed with short stiff pectinations. Proximal portion
of shaft covered dorsally by white and light carmine scales, remainder light carmine.
Head: Vertex white with a transverse carmine band mixed with sepia scales. Frons
bright yellow green (10GY 6/6 Munsell). Palpi vinaceous red laterally, white ventrally.
Thorax: Above bright yellow green (10GY 6/6 Munsell), shading to brownish red
at posterior margin, uniform creamy white below. Legs: fore and mid legs white,
speckled with carmine and vinaceous red on external surfaces; hind legs white. Ab-
domen: dorsal surface creamy white densely speckled with bright pink and sepia
scales. Segments 2, 3, 4 and 5 armed with prominent pink and white, laterally sepia
crests. Lateral and anal tufts creamy white. Venter creamy white. Upperside of fore-
wing: 13-18 mm from base to apex. Costa very slightly incurved near middle, apex
acute, but not falcate, outer margin slightly crenulate. Ground colour bright yellow
green (1O0GY 6/6 Munsell). Costa broadly white, densely speckled with bright pink
and sepia. An oval green spot at base of radius enclosed by an irregular whitish basal
area speckled with sepia and pink particularly at base and inner margin. Pale basal
area limited distally by narrow blackish antemedial and by some red scales. Ante
medial starts at costa at 1/7 from base, bends sharply proximad in base of cell, curves
regularly between cubitus and la, reaching inner margin at 1/4 from base. Stigma
reniform, very large, pale pink surrounded by a narrow dark ring, enclosing irregular
dark centre. Postdiscal very clearly defined, pale pink, narrowly edged on both sides
with mixed blackish and red scales, from costa at 3/4 from base, angled distad at
veins 7 and 6, broadening and bulging proximad from 6 to 4 narrowing and forming
a broad, even curve distad to centre of space 2 where it becomes the inner margin of
a large irregular pale tornal blotch. Tornal spot pale pink, narrowly edged with
blackish and red scales, enclosing some red, brown and green scales; proximal
margin of tornal spot (postmedial) curves abruptly proximad in space 1b reaching
inner margin at just over 1/2 from base. Distally edge of tornal spot is angled
distad at vein 2, then dips below it reaching termen at end of vein 2. A large
pinkish white marginal spot with irregularly crenulate proximal margin edged with
blackish and red scales, from apex to vein 3. Two small pinkish terminal lunules edged
proximally with blackish and red in spaces 2 and 3. Termen narrowly sepia, broadening
into small dark spots in spaces 4, 5, 6 and 7. Fringe whitish, mixed with some red and
blackish, particularly at end of veins. Hind wing: Outer margin regularly crenulated.
End of vein 6 especially produced. Ground colour as in fw, but costal area broadly
white. Some bright red scales at base and separating white costal area from remaining
green area. An elongated pinkish inner marginal area edged with blackish from base
to middle of inner margin. Stigma narrower than in fw and more densely obscured
by dark scales. Postmedial as in fw, of irregular thickness and shape but maintaining
approximately same distance from margin throughout. A very large black-edged
pinkish lunule in spaces 4—5 and a smaller one at tornus. A narrow blackish terminal

172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

line wider at middle of internervular spaces. Fringe as in fw. Underside: Very pale
green almost white with markings of upperside showing as pure white and faint red
terminal line widening into minute lunules between the veins.

Genitalia: Uncus short, pointed, slightly downcurved, base 2/3 of length with a
prominent lateral notch at each side. Subunci slender, membranous, hairy, shorter than
uncus. Gnathos directed ventrad, slightly constricted at base, terminating in an antero-
posteriorly flattened, bluntly pointed lobe armed with minute tubercles on posterior
surface. Saccus short, broad and almost truncated apically. Valve regular, long and
narrow, bluntly pointed apically. Harpe long and narrow, smooth, bluntly pointed
apically and curving inward with dorsal margin almost parallel to costa of valve, some-
what shorter than valve. Aedeagus: basal half long and much narrower than re-
mainder which is more lightly sclerotised except laterally and compressed into nu-
merous minute folds. Apical opening long, longitudinal, on ventral surface, armed at
base on left side by a blunt reflexed sclerotised lobe.

Holotype. ¢, Kakamega, Kenya, XII 1966, R. H. Carcasson and A. Forbes -Watson,
to be deposited in British Museum (Natural History). Paratypes: ¢@, Kakamega
Kenya XI 1967, R. H. Carcasson, to be deposited in British Museum (Natural History);
¢, all data as above, to be deposited in National Museum, Nairobi; ¢, locality as
above, III-1966, R. H. Carcasson, to be deposited in National Museum, Nairobi;
6, Mt. Elgon, Kenya, VII-1966, T. H. E. Jackson, to be deposited in National Mu-
seum, Nairobi.

This species is dedicated to my friend, the late Mr. T. H. E. Jackson of
Kitale, Kenya.

Archichlora sangoana Carcasson, new species
lines, @, is

Closely allied to A. pulveriplaga (Warren), (Figs. 8, 12), but differs in
having continuous, sinuous postmedial lines and smaller terminal blotches.

MALE. Antennae: Basal pectinations decreasing very gradually from base to apex.
Dorsal aspect of basal third of shaft covered by white scales. Head: Vertex white with
a reddish dot at base of each antenna and a reddish basal tuft at each side. Frons
bright green with some reddish scales at superior margin. Palpi whitish below, light
orange brown above. A fringe of reddish hair-scales at lower margin of eye. Thorax:
Uniform bright green above, except last segment which is light brown dorsally, yel-
lowish laterally. Ventral surface of thorax woolly, pale green anteriorly, white laterally
and posteriorly. Legs: Orange brown externally, creamy white internally. Abdomen:
Ist segment light brown with a small paler dorsal tuft. Subsequent segments very
pale brown with a dark dot flanking each side of the pale dorsal tufts. Underside uni-
form creamy white. Upperside of forewing: 17-20 mm from base to apex. Costa
very slightly concave near middle, evenly convex at 3/4 from base. Apex acute. Ter-
men moderately crenulate, tornus blunt. Ground colour green (25G 6/8 Munsell).
Costa creamy white, mixed with purplish and red scales at base. A narrow blackish
line separating white of costa from green ground colour near apex. Antemedial narrow,
greenisn yellow, from costa at 1/6 from base, to inner margin at 1/3 from base, slightly
angled distad at cubitus and at 1b. Postmedial narrow, greenish yellow, from costa
at 3/4 from base to inner margin at 2/3 from base, evenly and slightly curved distad
from costa to vein 3, more strongly curved proximad from vein 3 to inner margin.
Stigma large, pinkis white, surrounded by a ring of mixed blackish and red scales
with a central dot of the same colour. A large whitish terminal blotch from vein 7
almost to vein 4, the proximal edge of which projects proximad in space 6 and more
strongly so along vein 5. The pale blotch enclosing some bright ochreous scales and
a few blackish scales in spaces 5 and 4, and narrowly edged proximally by a mixture
of blackish and red scales. A similar but more rounded whitish blotch at tornus, in

VoLUME 25, NUMBER 3 Ws)

Figs. 10-13. Male genitalia of African Geometrinae. 10, Archichlora jacksoni sp.
nov., paratype; 11, Archichlora rectilineata sp. nov., paratype (Nairobi, Kenya); 12,
Archichlora pulveriplaga (Warren), (Kalinzu forest, Uganda); 13, Archichlora san-
goand sp. nov., paratype.

spaces la, lb and 2, with a narrow terminal extension in space 2. Termen narrowly
blackish, forming small internervular lunules in the areas occupied by the pale blotches.
Cilia whitish, chequered with blackish at the ends of the veins. Hind wing: Outer
margin somewhat produced at vein 6 and slightly concave between 6 and 4. Ground
colour as above. Postmedial as above, parallel to outer margin. Stigma very small.
A marginal blotch similar to those of fw but smaller and more regular between veins

174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

6 and 4, A very small whitish blotch at tornus, joined to a similar small triangle pointed
proximad along vein 1b, both edged with blackish and red scales. A small white patch
at base. Inner margin narrowly edged with pinkish, blackish and red scales and hairs.
Termen as in fw. Cilia uniformly grey, tipped with pinkish. Underside: Uniformly
white, with markings of upperside showing through very faintly. Termen and cilia
very narrowly blackish.

Genitalia: Uncus short, pointed, downcurved, almost as wide basally as long. Sub-
unci short, membranous, hairy, pointed, projecting laterad and posterad, almost as
long as uncus. Gnathos stout, pointed, projecting backwards, almost as long as uncus,
its inner (posterior) surface serrated. Saccus short, broad and blunt. Valve apically
pointed, tapering rapidly from a very broad base. Harpe broad, long, curved inwards,
irregularly shaped, with irregularly jagged margin, shorter than valve. A long sharply
pointed process projecting backwards from dorsal part of base of valve, armed dorsally
and on internal face with numerous sharp appressed teeth. Aedeagus long, basal half
much narrower than remainder, terminating in a pronounced rounded bulb-like ex-
pansion. Apical half membranous, compressed into numerous small longitudinal folds,
unarmed. Opening long, ventral, longitudinal.

Holotype. ¢, Katera, Sango Bay, Masaka, Uganda. X-1960, R. H. Carcasson.
Paratypes: 96 6, Malabigambo forest, Sango Bay, Uganda, II-1968, A. L. Archer.
Holotype and 4 paratypes to be deposited in British Museum (Natural History); re-
maining 5 paratypes to be deposited in National Museum, Nairobi.

Archichlora rectilineata Carcasson, new species
Wigs, 2..3)45.0, 9) LI
Superficially similar to A. pulveriplaga (Warren), (Figs. 8, 12), and
to A. sangoana Carcasson (Figs. 6, 13), but differs from both in being
smaller, more slenderly built and in having straight postmedial lines.

MALE. Antennae: Basal half of shaft decorated dorsally with white, black and
red scales. Head: Vertex white, sprinkled with red and black scales. Frons bright
green, palpi creamy, speckled dorsally and especially apically with rusty brown.
Thorax: Bright green (1OGY 5/6 Munsell) dorsally. Very pale green below, shading
to creamy white posteriorly. Legs: Very pale creamy buff, forelegs spotted with dark
brown on exterior surface. Abdomen: Above: 1st segment purplish brown, remaining
segments creamy buff, creamy white from segment 6 to apex. A pinkish dorsal crest
on first 4 segments. A dark brown dorsal spot on segments 5, 6 and 7. Below: uni-
formly creamy white. Upperside of forewing: 15-16 mm from base to apex. Costa
slightly but evenly arched. Apex acute. Outer margin slightly sinuous. Ends of veins
3, 4 and 7 slightly produced. Outer margin slightly concave from vein 4 to vein 7.
Ground colour bright green (1OGY 5/6 Munsell) with very faint paler vermiculations.
Costa whitish, densely speckled with blackish and red scales. A small patch of such
scales at base of costa. Antemedial narrow, light yellow-green, evenly curved from
costa at 1/5 from base to inner margin at 2/5 from base. Postmedial slightly wavy,
narrow, light yellow green, from costa at 3/4 from base to inner margin at 2/3 from
base. Stigma a smal] whitish dot edged with blackish and red scales. A very pale
pinkish, somewhat irregular, marginal lunule from just above vein 4 to middle of space
6, edged proximally by a very narrow blackish line. A similar but larger rounded
blotch at tornus enclosing an area speckled with dark scales. Terminal line blackish,
very narrow, of uniform width. Cilia whitish mixed with pink, blackish at the veins.
Hind wing: Margin deeply crenulate, veins 6 and 4 especially produced. Ground
colour as above. Postmedial as above, almost straight from just above junction of
veins 6 and 7 to inner margin just beyond 2/3 from base. Stigma very small, white.
A whitish marginal lunule similar to that of fw with its apex directed proximad along
vein 5. A similar but much smaller spot at tornus. Terminal line as in fw, cilia some-
what darker. Inner margin narrowly edged with blackish and red scales and by long

VOLUME 25, NUMBER 3 iS

Figs. 14, 15. Male and female genitalia of African Geometrinae. 14, Victoria
watsondé sp. nov., 6, paratype; 15, Archichlora rectilineata sp. nov., °, paratype.

whitish hairs. Underside: Generally whitish, shading to light green at costa and apex
of fw. Costa of fw narrowly blackish, stigma a minute reddish dot barely visible in
hw. Lunule of fw sepia, if present, with ill defined proximal edge. Terminal line and
cilia of both wings as above.

Genitalia: Genital capsule slender and lightly sclerotised. Uncus short, downcurved
and sharply pointed. Subunci broad, membranous, hairy, almost as long as uncus.
Gnathos a simple, bluntly pointed short projection. Saccus short, apically rounded.
Valve simple, regular, with apex gently pointed, rather broad at base, and with ventral
margin strongly elbowed midway from tip of saccus to apex. Harpe simple, broad,
close to ventral margin of valve, terminating in a smooth, inwardly curving spoon-like
lobe, apex of harpe not reaching apex of valve. Aedeagus lightly sclerotised, unarmed,
with basal bulb much shorter and broader than in other species. Opening of aedeagus
long, ventral.

176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

FEMALE. Similar to male, but larger, with more rounded wings. Fw 18-20 mm
from base to apex. Ovipositor lobes long and slender, curving inwards near apex. An-
terior and posterior struts slender and moderately long. Ostium simple, opening
posterad, vaginal plates very weakly sclerotised. Colliculum absent, ductus long and
slender with a sack-like expansion near base. Bursa long and slender, ovoid. Signum
at base of bursa, a smooth, large, irregularly rounded sclerotised plate with an inward
projecting, straight, transverse median ridge.

Two males from Mufindi, Iringa, Tanzania, belong to an individual
melanistic variant in which the dorsum of the abdomen is much darker,
the marginal lunules and blotches are completely filled in by blackish
scales and the cilia are uniformly blackish. A male from Kigezi, Western
Uganda, and a female from Njoro, Kenya are identical, with typical form,
but have a slightly curved postmedial in the hw and may represent a West
Rift subspecies.

Holotype. ¢ (pale form), Mufindi, Iringa, Tanzania, P. Burdon, to be deposited
in British Museum (Natural History); Allotype: 9 (pale form), Amani, Usambara,
Tanzania, X-1962, G. Pringle (No P 1188); Paratypes: 6 (pale form), Ngong,
Nairobi, Kenya, II-1954, Fowler and Coulson, in National Museum, Nairobi; ¢ (dark
form) Mufindi, Iringa, Tanzania, 1955, P. Burdon, to be deposited in British Museum
(Natural History); ¢ (dark form), data as above, in National Museum, Nairobi; 2
(pale form) Nairobi, Kenya, VI-1958, R. Carcasson, in National Museum, Nairobi.

Literature Cited

Carcasson, R. 1962. New African Lepidoptera. J. E. Afr. nat. Hist. Soc. 29 (105):
54-63.

Prout, L. B. 1912. Lepidoptera Heterocera, family Geometridae, sub-family Hemi-
theinae. Genera Insectorum, fas. 129.

1929-38. In Srrrz, A., The Macrolepidoptera of the World 16.

WarREN, W. 1898. New species and genera of the families Thyrididae, Uranidae,
Epiplemidae and Geometridae from the Old-World regions. Novit. zool. 5: 5-41.

BOOK REVIEW

A Key To THE RHOPALOCERA BUTTERFLIES OF Wyominc, by Clifford D. Ferris. Sci-
ence Monograph No. 21, Agriculture Experiment Station, University of Wyoming,
Laramie, Wyo. January 1971. 64 pp., 3 figs., 8 pls.

Dr. Ferris has tried to present in this slim volume a useable key to the butterflies
of Wyoming. Like all keys that I have seen, it is useful in the hands of those of us
who need no key and not so in the hands of a rank amateur. The eight plates at the
back of the book figure in black-and-white all of the species that the 4-H member
or beginner will find. The key to Colias and the discussions of Speyeria and Euphy-
dryas will be found helpful to those who have Wyoming material to name. An an-
notated checklist for the state is promised for “late 1971.” From what I have seen
of Ferris’s notes for this it is going to be a most useful index to the species found and
where they usually may be caught. Dr. Ferris did not see final page proof, so, there
are a few typographical errors that he would have caught.

F’, Martin Brown, Fountain Valley Rural Station, Colorado Springs, Colorado.

VOLUME 25, NUMBER 3 177

9HE LIFE HISTORY OF SCHINIA SEPARATA (NOCTUIDAE)

D. F. Harpwicx
Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario

Schinia separata (Grote, 1879 )1 is distributed in western North America
from southern British Columbia and Alberta southward to southern Cali-
fornia. In the Intermountain Basin area, it feeds in the larval stage on the
buds and blossoms of the sagebrush, Artemisia tridentata Nutt. (Figure
2). The moth is also common in some areas of the Great Plains in which
sagebrush does not occur but in which the hoary sagebush, Artemisia cana
Pursh. is abundant. Presumably in these grassland areas, the latter species
of Artemisia serves as food plant. The adult is a late-summer flier and is
present in most areas of its range during August and the first half of Sep-
tember. In the Intermountain Basin the period of adult activity is syn-
chronized with the budding period of the sagebrush and the moth flies
progressively later in the season with decreasing altitude and latitude.

Behaviour

Schinia separata is a nocturnal species, and the eggs are deposited in
the buds of Artemisia only at night. Three wild-caught females deposited
a mean of 91 eggs and the maximum deposited by a single female was
119. The behaviour of the ovipositing female separata was described in
my 1958 revision of the elliptoid-eyed Schinia:

“The female alights on a spray of the still tightly closed buds and crawls
over it, her abdomen arching ventrally. Having arrived at a site that is
apparently suitable, she secures herself by grasping buds and stems firmly
with clawed tarsi, and begins an elaborate exploration with the ovipositor.
The abdomen is arched ventrally and anteriorly, often being thrust for-
ward so as to protrude through the legs. The ovipositor, on encountering
the small hard bud, feels the surface and probes it with short, delicate
stabbing movements. When the edge of one of the stiff outer sepals of
the bud is encountered the blade of the knife-like ovipositor is forced
against this edge and the moth, by rapid lateral movements of the abdomen
which cause her whole body to vibrate inserts the ovipositor valves be-
neath the sepal. In this manner the egg is deposited within the bud at or
between the bases of the innermost sepals. The ovipositor is then with-
drawn and the moth usually flies to another bud cluster before the process
is repeated.”

1T cannot distinguish separata from Schinia acutilinea (Grt.) (1878, Can. Ent. 10: 232) with

any degree of certainty, but hesitate to synonymize the two without fuller study. If the two names
do represent the same species, acutilinea will take precedence.

178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1, 2. Schinia separata (Grt.) and its food plant. 1, Adult, Oliver, B. C.; 2,
Artemisia tridentata Nutt.

The egg is often grossly distorted in shape as a result of being inserted
beneath the sepals among the developing petals of the still tightly closed
bud. The majority of eggs hatch on the seventh day after deposition. The
newly hatched larva immediately attacks the succulent tissues of the de-
veloping bud. The larva usually completes its first stadium in the initial
bud, but during the second stadium migrates to an adjacent bud which it
enters by boring a hole through the apex where the sepals are thinnest.
The anterior two pairs of larval prolegs are reduced in size in the second
and third instars and the larva mixed with a looping motion similar to
that of catocaline larvae. During the third stadium the larva ceases to
conceal itself completely within the bud and feeds from an exposed po-
sition on the stem. The larvae continue to attack the buds at the apex,
however, where the tissues are softer.

Nineteen of 50 individually reared larvae matured in five stadia, the
remainder in six. The mature larva tunnels into the ground to form its
pupal cell and it is in the pupal stage that the insect remains throughout
the majority of the year.

Descriptions of Stages

The following descriptions of immature stages were based on the prog-
eny of three females taken in the Okanagan Valley near Oliver, B. C.
The larvae were reared individually at room temperature on the buds and
blossoms of Artemisia tridentata. Rearing techniques employed were those
used by Hardwick (1958). The estimate of variability following the
mean for various values is the standard deviation.

Unfortunately the larvae of separata being reared had to be removed
from the Okanagan Valley to Ottawa during the median stadia. Although
flowering heads of the food plant were shipped by air express to Ottawa,

VOLUME 25, NUMBER 3 179

the larvae did not develop normally and the ultimate stadium became
greatly protracted. The larvae eventually died without pupating. The
developmental data is therefore not complete, and the durations listed for
the median stadia may be somewhat attenuated.

Adult. (Fig. 1). Head and thorax silvery-fawn to greyish-fawn. Abdomen
silvery-fawn or silvery-grey. Underside of body dull pale grey. Forewing olivaceous
fawn marked with white; forewing occasionally with a rosy suffusion. Transverse
anterior line white, usually smooth, arched strongly and acutely outward. Basal space
olivaceous fawn, longitudinally streaked with white and with a dark margin along t.a.
line. Transverse posterior line white, generally smooth, excurved around cell, then
usually straight to inner margin. Median space fawn, paler than basal space, usually
with a pale longitudinal streak through cell. Orbicular spot not defined. Reniform
spot usually indicated only as a narrow dark shade. Subterminal line white, irregular,
closely approaching and occasionally fusing with outward arc of t.p. line. Subterminal
space darker than median space and usually darker than basal space. Terminal
space pale fawn. Fringe white checkered with fawn. Hind wing white with dark-
brown discal mark and dark-brown outer-marginal band. Outer-marginal band with
a white streak or blotch toward anal angle. Fringe white, usually with a pallid
brown inner line. Undersides of both wings white or pallid grey, marked with brown.
Forewing with small, dark-brown, orbicular and reniform spots, and with a paler brown
subterminal band. Hind wing with a narrow dark-brown discal spot and an evanescent
outer-subterminal band. Fringes of both wings white.

Expanse: 26.4 + 2.2 mm (92 specimens).

Egg. Very pale greenish-yellow when deposited. Losing some of greenish colouring
on the second day, then remaining essentially unchanged for remainder of incubation
period. Mandibles and ocelli becoming visible through chorion a few hours before
hatching.

Incubation period: 7.2+0.7 days (157 eggs).

First-Stadium Larva. Head translucent blackish-grey. Prothoracic and suranal
shields somewhat paler than head. Trunk pale greyish-white becoming stained with
yellow or green after larva begins to feed. Thoracic legs and proleg shields dark
smoky-brown.

Head width: 0.240 + 0.011 mm (25 larvae).

Duration of stadium: larvae maturing in five stadia, 5.3 + 0.7 days (19 larvae);
larvae maturing in six stadia, 5.1 + 0.4 days (31 larvae).

Second-Stadium Larva. Head varying from dark brown to medium light brown.
Prothoracic shield concolorous with trunk except for some pale-brown flecking.
Suranal shield concolorous with trunk. Trunk with a chalky opacity uncommon in
second-stadium noctuid larvae; varying from a blotchy yellowish-grey to a blotchy
greenish-grey. Thoracic legs and proleg shields brown. First two pairs of anal prolegs
smaller than remaining pairs.

Head width: 0.319 + 0.010 mm (25 larvae).

Duration of stadium: larvae maturing in five stadia, 4.0 + 0.6 days (19 larvae);
larvae maturing in six stadia, 4.1 + 0.6 days (31 larvae).

Third-Stadium Larva. Head pale fawn grey. Prothoracic shield whitish-grey
marked with brown. Suranal shield essentially undistinguished from trunk. Mid-
dorsal area dull, dark brownish-grey. Subdorsal area white or pale grey with a pair
of dark-grey median lines. Supraspiracular area dark greyish-brown, with a discon-
tinuous, white or pale-grey median line. Spiracular band white or pale grey with a
darker-grey or brown median shade. Suprapodal area similar in colour to supra-
spiracular area. Mid-ventral area varying from greenish-grey to brownish-yellow.

Head width: 0.51 + 0.02 mm (25 larvae).

Duration of stadium: larvae maturing in five stadia, 6.1 + 1.2 days (19 larvae);
larvae maturing in six stadia, 4.9 + 1.1 days (31 larvae).

180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Antepenultimate-Fourth-Stadium Larva. Head fawn-grey heavily marked with
medium brown. Prothoracic shield white, usually heavily suffused with brown; heavily
suffused shields with three well-defined longitudinal white lines. Suranal shield
concolorous with trunk. Mid-dorsal band dark-brown with a pale median shade. Sub-
dorsal area pale grey or white with a pair of pale-brown median lines. Supraspiracular
area brown with a badly broken white or pale-grey median line. Spiracular band white
with an irregular light-brown median line. Suprapodal area concolorous with supra-
spiracular area. Mid-ventral area pale grey mottled with light brown.

Head width: 0.74+ 0.03 mm (44 larvae).

Duration of stadium: 5.7+ 1.1 days (31 larvae).

Penultimate-Stadium Larva. Head creamy white mottled with mauve-brown.
Prothoracic shield white with a broad median and a pair of narrower submarginal lines
of mauve-brown. Suranal shield poorly distinguished from remainder of trunk. Mid-
dorsal band dark brown with a pale median shade. Subdorsal area white with a pair
of irregular, pale-mauve median lines. Supraspiracular area mauve-brown, paler dor-
sally than ventrally; mottled mesally with white. Spiracular band white with an ir-
regular and discontinuous, mauve median band. Suprapodal area concolorous with
supraspiracular area. Mid-ventral area grey variably suffused with mauve.

Head width: fourth-stadium larvae maturing in five stadia, 0.95 + 0.06 mm (30
larvae); fifth-stadium larvae maturing in six stadia, 1.08 + 0.04 mm (37 larvae).

Duration of stadium: fourth stadium of larvae maturing in five stadia, 7.6 + 1.0
days (19 larvae); fifth stadium of larvae maturing in six stadia, 6.4+1.0 days (31
larvae ).

Ultimate-Stadium Larva. Head whitish-grey, heavily mottled with medium brown.
Prothoracic shield white suffused and mottled with light brown and with a median
and a pair of subdorsal lines devoid of darker shading. Suranal shield essentially un-
distinguished from remainder of trunk. Mid-dorsal band dull dark brown, with a var-
iably expressed, paler, mauve median shade. Subdorsal area white marked with
purplish-brown mesally and with a pair of mauve, median longitudinal lines. Supra-
spiracular area dark brown irregularly marked with mauve and cream. Spiracular
band creamy-white with a discontinuous and irregular, mauve median line. Supra-
podal area purplish-brown marked with cream. Mid-ventral area grey variably suffused
with purplish-brown.

Head width: 1.75+0.08 mm (25 larvae).

Acknowledgment

I appreciate the assistance of my associate, Mr. Eric Rockburne, in
measuring the head widths of the larvae.

Literature Cited

Grote, A. R. 1879. New noctuids. Can. Ent. 11: 197-199.

Harpwickx, D. F. 1958. Taxonomy, life history and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae) with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

VoLuME 25, NUMBER 3 181

THE LIFE HISTORY OF SCHINIA WALSINGHAMI (NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute, Canada Department of Agriculture,
Ottawa, Ontario

Schinia walsinghami (Hy. Edwards, 1881) feeds in the tarval stage on
rabbit brush, Chrysothamnus nauseosus Nutt. (Fig. 3). The moth is dis-
tributed in the Intermountain Basin area of western North America from
the Okanagan Valley of British Columbia southward to southern Cali-
fornia. Records available in the Canadian National Collection (Helena,
Montana; Arvada and Alcova, Wyoming) suggest that the species occurs
only marginally on the Great Plains, although the food plant itself is much
more widely distributed.

The flight period of walsinghami is closely co-ordinated with the late-
summer blossoming period of Chrysothamnus, the majority of specimens
examined having been taken in the latter half of August and during the
month ot September. The species is single-brooded throughout its range.

Behaviour

The adult of Schinia walsinghami is nocturnal and the eggs are de-
posited only during the hours of darkness. The oviposition pattern is
similar to that of Schinia separata (Grt.) and the female works vigorously
to deposit its eggs among the developing florets in the small hard buds of
the rabbit brush. The ovipositor may be inserted through the top of a
partially opened bud or between the stiff sepals forming the lateral walls
of the bud. As with separata, the eggs are often badly distorted as a result
of being wedged among the florets. Of two individually confined wild-
caught females, one deposited a total of 69 eggs and the other 65. The
majority of eggs hatched on the sixth day after deposition.

The newly hatched larva commonly bores into a floret and tunnels
downward through it until it reaches the seed, then crosses over into an
adjacent floret and tunnels upward through this. Usually early in the
third stadium the larva leaves the first bud and attacks a second which it
enters through the tip. By the time that it is in the late third stadium, the
larva is becoming rather large to accommodate itself readily in the Chryso-
thamnus bud and often the caudal end may be found protruding from the
apex of the bud. Fourth and fifth instars do not conceal themselves within
individual buds but feed on them from a position on the stem, the larva
reaching the florets by boring a hole through the sepals at the side of the

bud.

All the individually reared larvae matured in five stadia. At the ces-

182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 1-5. Schinia walsinghami Hy. Edw. and its food plant. 1, Adult, Biggs, Oreg.;
2, 5, dorsal aspect of ultimate-stadium larvae; 3, food plant, Chrysothamnus nauseosus
Nutt.; 4, lateral aspect of ultimate-stadium larvae.

VOLUME 25, NUMBER 3 183

sation of feeding the larva tunnels into the ground to form its pupal cell
and it is in the pupal stage that the species overwinters.

Descriptions of Stages

The following descriptions of immature stages are based on the progeny
of two females taken near Oliver in the Okanagan Valley of British Co-
lumbia. The larvae were reared individually at room temperature using
methods outlined by Hardwick (1958). The estimate of variability fol-
lowing the mean for various values is the standard deviation.

Adult (Fig. 1). Head and thorax pale fawn-grey. Abdomen silvery-grey. Under-
side of body pale silvery-grey. Forewing light reddish-brown to golden fawn variably
marked with white. Transverse posterior line white, strongly and usually smoothly
excurved, sometimes weakly denticulate. Basal space reddish-brown to fawn, darker
outwardly than at base, often with longitudinal pale streaking. Transverse posterior
line white, bisinuate, excurved around cell, then incurved to trailing margin, usually
strongly denticulate. Median space paler brown than basal space, rather diffusely
marked and sometimes with some pale longitudinal streaking; median space often paler
on either side so that white of t.a. and t.p. lines obscured; orbicular spot not defined
and reniform spot usually indicated only as a dark shade. Subterminal line irregular,
usually indented opposite cell and toward trailing margin. Subterminal space con-
colorous with basal space. Terminal space cream, variably suffused with light fawn;
often a series of intervenal brown points at margin of wing. Fringe cream or pale
fawn, often checkered with brown. Hind wing white or cream, variably suffused
with brown; with a broad, light-brown outer-marginal band and a light-brown discal
lunule. Outer-marginal band usually with a pale median streak. Fringe white, often
with a brown median line. Underside of both wings shining greyish-cream, each with
an. indistinct, light-brown, discal spot and post-median band. Fringe of forewing
cream, often checkered with brown; fringe of hind wing white.

Expanse: 24.2 + 1.6 mm (100 specimens ).

Egg. Pale greenish-yellow when deposited. Losing greenish colouring on day after
deposition, then gradually becoming a somewhat darker yellow during the next few
days. Mandibles, then ocelli, and then the whole head capsule becoming visible
through chorion on the day of hatching.

Dimensions of egg: length, 0.791 + 0.043 mm; diameter, 0.413 + 0.023 mm
(20 eggs).

Incubation period: 6.00+0.2 days (63 eggs).

First-Stadium Larva. Head medium to dark brown. Prothoracic and suranal
shields somewhat lighter brown. Trunk creamy-white on hatching but becoming
stained with yellow after feeding.

Head width: 0.285 + 0.008 mm (25 larvae).

Duration of stadium: 4.6 + 0.5 days (20 larvae).

Second-Stadium Larva. Head medium brown. Prothoracic and suranal shields
lighter brown. Trunk greyish-white on entering stadium, becoming light yellow after
feeding.

Head width: 0.440 +0.019 mm (25 larvae).

Duration of stadium: 3.6 + 0.5 days (20 larvae).

Third-Stadium Larva. Head orange-brown. Prothoracic shield orange-yellow with
some light-brown mottling. Suranal shield essentially undistinguished from trunk.
Mid-dorsal band orange-brown, narrow. Subdorsal area greyish-yellow margined
laterally by a brown line. Supraspiracular area usually darker than, occasionally con-
colorous with, subdorsal area. Spiracular band dull white. Suprapodal area usually
paler than supraspiracular area. Mid-ventral area pale greyish-yellow.

184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 6, 7. Schinia walsinghami Hy. Edw., apical abdominal segments of pupa. 6,
Ventral; 7, right lateral.

Head width: 0.694 + 0.019 mm (25 larvae).

Duration of stadium: 6.1 + 0.7 days (20 larvae).

Fourth-Stadium Larva. Head orange-brown. Prothoracic shield paler orange-
brown; variably marked with medium brown, most strongly so near margins. Suranal
shield poorly distinguished from trunk, usually with some brown stippling. Mid-dorsal
band orange-brown, the darkest area of trunk. Subdorsal area fawn, margined laterally
by a pale-grey and a dark-brown lines. Supraspiracular area concolorous with sub-
dorsal area. Spiracular band narrow, dull grey, poorly defined. Ventral region
yellowish-grey.

Head width: 1.13 + 0.04 mm (25 larvae).

Duration of stadium: 7.5 + 0.5 days (20 larvae).

Fifth-Stadium Larva (Figs. 2, 4, 5). Head medium to dark orange-brown.
Prothoracic shield paler, variably mottled with brown, most strongly so along lateral
and posterior margins; a paler mid-dorsal line usually evident on shield. Suranal shield
pale orange, variably stippled with brown. Trunk varying from translucent pale
orange to translucent yellowish-grey; maculation of trunk usually indistinct. Mid-
dorsal band orange-brown, narrow. Subdorsal area greyish-yellow. Supraspiracular
area somewhat darker yellow than subdorsal area. Spiracular band reduced to a poorly
defined, narrow pale shade. Ventral region pale greyish-yellow.

Head width: 1.73 + 0.04 mm (9 larvae).

Duration of feeding phase of fifth stadium: 10.3 + 2.0 days (20 larvae).

Duration of prepupal phase of fifth stadium: 5.6 + 1.4 days (20 larvae).

Pupa (Figs. 6, 7). Pale yellowish-brown. Spiracles on abdominal segments 4
to 7 sunk into deep pits. Anterior margins of segments 5, 6 and 7 each with a broad
band of prominent pitting. Proboscis terminating between apex of wings. Cremaster
consisting of two slender setae borne on a short rounded prolongation of the tenth
abdominal segment; a pair of vestigial lateral setae also usually evident.

Length from anterior end to posterior margin of fourth abdominal segment: 6.8 +
0.2 mm (3 pupae).

Acknowledgments

I am grateful to Mr. John E. H. Martin of the Entomology Research In-
stitute for the photographs accompanying this paper, and to my associate

VoLUME 25, NUMBER 3 185

Mr. E. W. Rockburne for measuring the immature stages and for drawing
pupal structures.

Literature Cited

Epwarps, H. 1881. New genera and species of North American Noctuidae. Papilio
1: 19-28.

Harpwick, D. F. 1958. ‘Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

DISTRIBUTION AND GEOGRAPHICAL DIFFERENTIATION
OF MARPESIA ELEUCHEA HUBNER (NYMPHALIDAE), WITH
DESCRIPTIONS OF TWO NEW SUBSPECIES

EUGENE MUNROE

Entomology Research Institute
Canada Department of Agriculture, Ottawa, Ontario

The tailed butterfly Marpesia eleuchea Hiibner appears, despite some
doubtful continental records, to be an Antillean endemic. Records from
the United States are based either on misidentified specimens of M.
petreus (Cramer) (e.g., Slosson records from Miami) or on specimens of
very doubtful provenance (e.g. the Doll specimen from “Central Florida”
mentioned by Kimball (1965) and the “Southern States” specimen men-
tioned in the present paper). A record from Colombia discussed below is
also highly suspect. The two species M. eleuchea and M. petreus are
recognized by almost all authorities (e.g., dos Passos, 1964) as being dis-
tinct. Hemming’s statement (1967, p. 277) that the two are currently
treated subjectively as subspecies appears to be based on outdated in-
formation. Allowing for the wrong or questionable records noted above,
the two are almost completely allopatric. M. eleuchea inhabits Cuba,
Hispaniola, Jamaica and the Bahamas, whereas M. petreus has a main-
land range extending from the southern United States far into South
America, and also from Trinidad up the Lesser Antillean chain into the
Virgin Islands and Puerto Rico, where M. eleuchea might have been
expected but appears not to occur. There is a possible minor area of
overlap in the Bahamas, where M. eleuchea occurs on New Providence
and possibly other islands, while M. petreus is represented by a specimen
in the British Museum (Natural History) labelled, “Bahamas, July 1921,
J. M. St. J. Yates”. However, it is not certain that the two species occur
together on the same islands in the Bahama group.

186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

This pattern of geographical replacement is of some evolutionary in-
terest. The endemic West Indian M. eleuchea is the most closely similar
species to M. petreus, but is manifestly more primitive; for example the
shape of the wings and the configuration of the forewing bands are less
divergent from those of more normal nymphalids, and the ocellate spots
are less degenerate. It is reasonable to suppose that it is a West Indian
stage of evolution of the stock which on the mainland has developed into
the more strongly apomorphic M. petreus. The latter, however, has been
successful in invading the West Indies via the Lesser Antilles. These
small islands do not have a significant endemic butterfly fauna, but are
populated by species of three geographical categories. First there are
widespread species, some of which, like Battus polydamas (Linnaeus),
have developed endemic subspecies and some of which, like Ascia
monuste (Linnaeus), have not. Second, there are Greater Antillean species
which have spread southward down part of the Lesser Antillean chain.
An example is Heliconius charitonius (Linnaeus ), whose subspecies reach
St. Kitts. Third, there are South American species which have moved
northward into the lesser Antilles, usually without differentiation, like
Biblis hyperia (Cramer), but occasionally with minor subspeciation, as
in Mestra cana (Erichson). M. petreus is an extreme example of the
latter type. Not only has it occupied the whole of the Lesser Antilles but it
has spread through the Virgin Islands and Puerto Rico as well. When
Antillean endemics that occur in Hispaniola are represented in Puerto
Rico as well, the two populations are usually identical or at least very
similar. It is most ususual for an unmodified continental vicariant to
occur in Puerto Rico instead, as in the present case. It seems very likely
that Marpesia eleuchea originally inhabited Puerto Rico (and perhaps
the Virgin Islands) as well as the other Greater Antilles, and that it has
been displaced by competitive exclusion following the arrival of M.
petreus from the south. An alternative hypothesis is that M. eleuchea
never occurred in Puerto Rico or died out there before the arrival of M.
petreus, which penetrated this “empty” habitat but has so far failed to
establish itself sympatrically with M. eleuchea in Hispaniola.

It has been known for a long time that M. eleuchea has significant
geographical variation in the Antilles. Bates (1935, p. 172), for example,
said of M. eleuchea, “This choromorph is limited to Cuba and the Ba-
hamas; specimens from Hispaniola and Jamaica (pellenis Godt) are less
heavily marked.” Some years ago when I was studying the biogeography
of West Indian butterflies, the late Mr. William P. Comstock called my
attention to the geographical variation of this species and indicated that
Mr. C. F. dos Passos intended to work it out. At Mr. Comstock’s sugges-

VoLUME 25, NUMBER 3 187

Figs. 1-4. Marpesia eleuchéa eleuchea Hiibner. 1, 2, 6, Upper and under sides,
Santiago de Cuba, F. E. Church, AMNH;; 3, 4, 2, upper and under sides, Santiago de
Cuba, F. E. Church, AMNH.

tion, Mr. dos Passos kindly relinquished study of the problem to me and
I set aside a type series of a manuscript subspecies from Hispaniola. The
present paper describes that subspecies and another from the Bahamas,
and compares these with the two named subspecies.

Marpesia eleuchea eleuchea Hibner
Figs. 1-4.

Marpesia eleuchea Hiibner, 1818, p. 32, pl. [35], figs. 197, 198.

Marpesia eleuchea: Hibner, 1819, p. 47.

Marpesia eleuchea: Hiibner [1823], pl. [50].

Nymphalis eleuchea: Poey, 1847, p. 47.

Marpesia eleucha: Doubleday, 1844, p. 86. In part.

Timetes (Marpesia) eleucha: Doubleday, Westwood and Hewitson, 1850, p. 263. In
part.

Timetes eleuchea: Herrich-Schaffer, 1864, p. 161.

Megalura eleucha: Gundlach, 1881, p. 112.

188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Athena eleucha: Dyar, 1902, p. 25. In part.

Megalura eleuchea: Seitz, 1914, p. 470.

Athena eleuchea eleuchea: Bates, 1935, p. 172. In part.

Marpesia eleuchea: Comstock, 1944, p. 462. In part.

Marpesia eleucha: dos Passos, 1964, p. 74. In part.

Marpesia eleuchea: Hemming, 1967, p. 277. In syn. M. petreus (Cramer).

Male. Forewing short, termen erect and with moderately produced subapical
angle. Upperside fulvous, with three oblique black bars in cell, the second broken,
the third produced to anal vein. A black bar on discocellular. Three black lines
between cell and termen: the first oblique to Ms, there angled and erect to posterior
margin; the second narrow, anteriorly faint and weakly curved; the third thick, a
little diffuse, parallel to termen. Apical area infuscated.

Hind wing above similar in colour. A black line from costa to Cu, traversing cell.
A bent, broken, black bar at end of cell. A narrow black postmedial line, erect from
costa to Cu», there curving basad and fading out. Three subterminal bands. The first
fuscous, diffuse, preceded on costa by a white shade and followed by a greyish
apical shade, weakly converging with postmedial to M:, there bent basad and scal-
loped to form basal edges of three large, grey, suffused, ocellate spots. The second
subterminal band diffuse, fuscous, converging with the first from near apex to M:,
thence scalloped to form distal borders of ocellate spots. The third subterminal band
narrower, better defined, close to and parallel to termen, forming a deep dentation
into the tail. Anal area suffused with greyish fuscous, the suffusion not taking in
all of the first ocellate spot. Tail white-tipped. Anal lobe blue-suffused.

Underside brown with violet tints, paler beyond middle, leaf-like. Forewing with
first cell-band reduced, curved, brown. Second cell-band broken in middle. Third
cell-band narrow, irregular, black, distally pale-edged. Discocellular bar grey, edged
with fuscous powdering. Postmedial line light grey, followed by dark brown, form-
ing the midrib of the leaf-like pattern. A complete series of poorly defined ocellate
spots, bordered by weak, scalloped, brown, first and second subterminal lines. Third
subterminal line following same course as on upperside, but very weak.

Hind wing beneath coloured like forewing. Pattern elements occupying same posi-
tions as above, but with a complete series of ocellate spots.

Female. Upperside like that of male, but much duller fulvous, lines thicker. Fore-
wing and hind wing with suggestions of complete series of ocellate spots, defined
outwardly by greyish-fuscous suffusion. Whole terminal area suffused with greyish
or brownish fuscous. Apical suffusion of forewing brownish fuscous, not black.

Underside like that of male, but grey rather than brown.

ee The type is, so far as I know, lost, but Hiibner gives the locality Havana,
Cuba.

Material examined. Eighty-three specimens, from Cuba: Rangel, P. del R:;
Havana, June; Nueva Gerona, I. of Pines, Aug.; Santiago de las Vegas, July; Cama-
guey, April; Soledad, Sta. Clara, July; Matanzas, July; Santiago de Cuba, 200 ft.,
June; Torquino R., 100 ft. July; Sierra Maestra, 1000 ft., March, June; Loma del Gato,
Sierra Maestra, 850 m., July; Guantanamo, June; Baracoa; Holguin; Rio Cano, March;
Tanamo, March; San Christobal. Two specimens labelled Guacomo, Colombia, F.
EK. Church, appear to belong to this subspecies.

Marpesia eleuchea bahamensis Munroe, new subspecies
Figs. 5-8

Athena eleuchea eleuchea: Bates, 1935, p. 172. In part.

Male. As in the nominate subspecies, with the following differences. Size a little
smaller on average. Markings of forewing slightly heavier. Hind wing above with
apex grey-suffused from first subterminal band to termen. Second subterminal band

VOLUME 25, NUMBER 3 189

Figs. 5-8. Marpesia eleuchea bahamensis n. subsp. 5, 6, Holotype, ¢, upper and
under sides, Nassau, Bahamas, F. E. Taylor, BM (NH). 7, 8, allotype, 2, upper and
under sides, Nassau, Bahamas F. E. Taylor BM(NH).

diffuse, with fuscous suffusion extending weakly distad to termen. Third subterminal
line heavier. Anal area with greyish-fuscous suffusion more extensive, completely
encompassing first ocellate spot as well as second and third. Basal margins of ocellate
spots each with a strong white lunular marking. Some light-grey suffusion in outer
parts of ocellate spots and in some specimen also along third subterminal band
posteriad of tail.

Underside purplish grey, with weak markings.

Female. Larger and duller than male. Like female of the nominate subspecies,
but with bands of forewing above heavier. Hind wing with more fuscous and grey
sutfusion and stronger white anal markings, as in male.

Underside light grey, with weak markings.

Types. Holotype, male, and allotype, female, Nassau, Bahamas, F. E. Taylor, In
British Museum (Natural History). Three male and 3 female paratypes in the
British Museum (Natural History) from the Bahamas: [Andros?], N. Chamberlain;
Nassau, J. L. Bonhote, F. E. Taylor; “Bahamas, 10.6.1900”, Sir G. Carter. Four
additional paratypes in the Museum of Comparative Zoology, Cambridge, Mass.,
from the Bahamas: Nassau, June; Andros, July; Arthur's Town, Cat. I., July. In the
Carnegie Museum one paratype Blue Hills, Nassau, Jan.

190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 9-12. Marpesia eleuchea dospassosi n. subsp. 9, 10, Holotype, ¢, upper
and under sides, Punta Arena, San Lorenzo, Rep. Dominicana, 24 June 1915, F3177.
AMNH; 11, 12, allotype, 2, upper and under sides, Pétionville, Haiti, 12 June 1930,
F,. E. Church, AMNH.

Marpesia eleuchea dospassosi Munroe, new subspecies
Figs. 9-12
Marpesia eleuchea pellenis: Bates, 1935, p. 172. In part.

Male. Coloured like the nominate subspecies. Forewing above with subapical
angle on the average more rounded and termen more shallowly curved and oblique.
Postcellular continuation of the third cell-stripe tending to be displaced basad. An-
terior part of postmedial band strongly thickened towards costa. First subterminal
line faint posteriorly, nearly obsolete anteriorly. Apical black suffusion wide, con-
tinued around apex to join expanded outer subterminal band. The latter wide, black
and reaching nearly to termen.

Hind wing above with discocellular bar tending to be weak. First postmedial line
usually narrow and broken. Second postmedial line nearly obsolete anterior to Mb.
Ocellate spot in cell Mz largely fulvous.

Wings below as in nominate subspecies but variable in colour, from grey through
tan to dark brown.

Female. Upperside closely similar to that of the nominate subspecies, but with

VoLUME 25, NUMBER 3 191

terminal area more broadly and heavily suffused with brownish fuscous on forewing
and hind wing.

Underside as in nominate subspecies, but with ocellate spots more distinct.

Types. Holotype, male, Punta Arena, San Lorenzo, Dominican Republic, 24 June
1915, F3177, in the American Museum of Natural History. Allotype, female, Pétion-
ville, Haiti, 12 June 1930, F. E. Church, in the American Museum of Natural History.
Five male, 3 female paratypes in the American Museum of Natural History, same data
as holotype, and: La Romana, Dominican Republic, 14 Dec., G. N. Wolcott; Rio
Macosia, 8 km. south of Las Matas, Dominican Republic, 5 July 1957, O. Cucurullo
Jr.; Pétionville, Haiti, 3 June 1930, F. E. Church; Cit. La Ferriére, Haiti, 2,500 ft.,
6 June 1935. Five male, 4 female paratypes in the British Museum (Natural His-
tory): “Haiti”, Tweedie; Port-au-Prince, Haiti, F. Odile Joseph; “Gulf of Mexico,
P. E. Cheesman”; [Haiti], ex coll. Chris. Ward. Five paratypes in the Cornell Uni-
versity Collection: Pétionville, Haiti, May, June; Monte Christi, Dominican Republic,
June. Three paratypes in the Museum of Comparative Zoology, Cambridge, Mass.:
“San Domingo”.

Marpesia eleuchea pellenis (Godart)
Figs. 13-16

Nymphalis pellenis Godart, 1819, p. 359.

Marpesia pellenis: Doubleday, 1844, p. 86. In syn. of M. eleucha.

Timetes (Marpesia) pellenis: Doubleday, Westwood and Hewitson, 1850, p. 263.
In syn. of T. eleucha.

Megalura eleucha: Kaye, 1931, p. 533. In part.

Athena eleuchea pellenis: Bates, 1935, p. 172. In part.

Marpesia eleuchea: Comstock, 1944, p. 462. In part.

Marpesia eleuchea pellenis: Avinoff and Shoumatoff, 1946, p. 280.

Male. Like M. e. dospassosi in colour. Forewing with subapical angle more acute
and termen more strongly excavated than in other subspecies. Transverse lines finer.
Postcellular continuation of third cell line displaced basad, very fine. Postmedial line
moderately expanded towards costa. First subterminal line very weak, especially
anteriorly. Third subterminal narrow, removed from termen, interrupted or nearly
so between M; and M>. Apical patch extending to subapical angle, its posterior mar-
gin nearly straight, oblique.

Hing wing above with discocellular line weak or absent. Postmedial and first sub-
terminal lines narrower than in M. e. dospassosi. First ocellate spot almost wholly
fulvous. Third subterminal line narrow.

Underside as in M. e. dospassosi.

Female. Smaller and paler than the female of M. e. dospassosi. Transverse lines
narrower than in other subspecies. Costal, apical and terminal suffusion of forewing
and hind wing greatly reduced.

Underside as in M. e. dospassosi.

Type. Godart described pellenis from a specimen without exact locality, of which
he said, “Nous la soupconnons des Antilles.” The material studied by Godart came
from several collections, and though I think it doubtful that his type still exists it is
possible it may yet be discovered. I therefore refrain from designating a neotype.
His description refers to the absence of the third black line, i.e., the first subterminal,
which would point to either this subspecies or the preceding one, but does not give
any hint as to which of the two Godart had. I consider that he is as likely to have
received a Jamaican specimen as one from Haiti, and I therefore arbitrarily restrict
the type locality to Jamaica, making the name applicable to the present subspecies.

Material Examined. Thirty specimens from Jamaica: Montego Bay, May; Baron
Hill, Trelawny, 1,150 ft., Aug., Sept., Oct., Nov.; Reading, St. James; Hope Botanic

192 JoURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 13-16. Marpesia eleuchea pellenis (Godart). 13, 14, Male, upper and
under sides, Hope Botanic Gardens, St. Andrew, Jamaica, 5 Dec. 1919, F4455,
AMNH; 15, 16, 2, upper and under sides, Baron Hill, Trelawny, Jamaica, Noy. 1935,
AMNH.

Gardens, St. Andrews; Fish River, Portland; Constant Spring, 650 ft.; Rae Town,
Aug.; Albion, July; Milk River, June. Also one specimen without locality, and one
labelled “Southern States, No. 1442, Coll. J. Angus, West Farms, New York.”

Acknowledgments

I wish to express appreciation to Mr. C. F. dos Passos and the late Mr.
William P. Comstock for encouraging me to undertake this study. The
late Mr. F. E. Watson, the late Dr. A. Avinoff, the late Dr. Wm. T. M.
Forbes, Professor V. Nabokov, Dr. P. J. Darlington Jr. and Dr. F. H.
tindge lent material or helped in other ways. Mr. T. G. Howarth helped
me to examine material at the British Museum and kindly provided
photographs of the types of Marpesia eleuchea bahamensis. Messrs. Orrin
Manright and D. Kritsch made the remaining photographs and Mr. Kritsch

VoLUME 25, NUMBER 3 193

mounted the plates. Mr. E. C. Pelham-Clinton gave prompt and helpful
information on specimens in the Dufresne Collection in the Royal Scottish
Museum, Edinburgh, and Dr. Pierre Viette verified the absence of type
material in the Muséum d Histoire Naturelle, Paris.

Literature Cited

AvINoFFr, A., AND N. SHOUMATOFF. 1946. An annotated list of the butterflies of
Jamaica. Ann. Carnegie Mus. 30: 262-295.

Bates, M. 1935. The butterflies of Cuba. Bull. Mus. Comp. Zool. 78: 63-258.

Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands. Lepidoptera
Rhopalocera. New York Acad. Sci., Sci. Surv. Porto Rico and the Virgin Is.
12: 419-622.

pos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lep. Soc.
Mem. l.

DovusLepay E. 1844. List of the specimens of lepidopterous insects in the collec-
tion of the British Museum. Part 1. London.

Dousiepay, E., J. O. WEstwoop and W. C. Hewirson. 1850. The genera of
diurnal Lepidoptera. Signature 4A. London.

Dyar, H. G. 1902. A list of North American Lepidoptera. U.S. Natl. Mus. Bull.
52-8

Gopart, J. B. 1819. In Latreille and Godart, Papillon, Encyc. Méth., Hist. Nat.
9. Paris.

GunpiAcH, J. 1881. Contribucién 4 la entomologia cubana. I. Lepidoptera.
Havana.
Hemminc, F. 1967. The generic names of the butterflies and their type-species
(Lepidoptera: Rhopalocera). Bull. Brit. Mus. (Nat. Hist.), Ent. Suppl. 9.
HeErRRICH-SCHAFFER, G. A. W. 1864. Die Schmetterlingsfauna der Insel Cuba.
Correspbl. Zool.-Min. Ver. Regensburg 18: 159-172.

HoLianp, W. J. 1916. The Lepidoptera of the Isle of Pines [etc.] Ann. Carnegie
Mus., 10: 487-518, pl. 31.

Hupsner, J. [1818] Zutrage zur Sammlung exotscher Schmettlinge. Vol. 1, text,
validating Vol. 1, pl. [35], [1814-18].

Husner, J. [1819]. Verzeichniss bekannter Schmettlinge. Signature 3, Augsburg.

Hutsner, J. [1823]. Sammlung exotischer Schmetterlinge. 2, pl. [50]. Augsburg.

Kaye, W. J. 1931. Additions and corrections to the author’s “Butterflies of Jamaica
(1926).” Trans. Ent. Soc. London 79: 531-537, pl. 39.

Kimpa.1, C. P. 1965. The Lepidoptera of Florida, an annotated checklist. Gaines-
ville.

Pory, Pu. 1947. Catalogo metédico y descriptivo de las mariposas de la isla de
Cuba. Mem. Real Soc. Economica, Habana 3: 44-50.

Serrz, A. 1907-1924. The Macrolepidoptera of the World. V. The American
Rhopalocera. Stuttgart.

194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

SEVEN NEW LEPIDOPTEROUS LEAF-MINERS
ASSOCIATED WITH QUERCUS AGRIFOLIA
(HELIOZELIDAE, GRACILARIIDAE )

PauL A. OPLER

Division of Entomology
University of California, Berkeley

During the course of a study of the bionomics of Microlepidoptera asso-
ciated with Quercus agrifolia Neé (Fagaceae) in California, it was found
that a number of the species are undescribed. In order to provide names
for use elsewhere, one new heliozelid and six new gracilariids are herein
described. Because the genitalia of the Nearctic members of these fam-
ilies have not been previously studied, the works of Kumata (1963),
Kuroko (1961), and Vari (1961) provided bases of homologies and diag-
nostic features in the genitalia of these moths.

Descriptions in this paper use the holotype and allotype as exemplars.
The holotype and allotype of each species are deposited in the California
Academy of Sciences, San Francisco, on indefinite loan from the Califor-
nia Insect Survey, University of California, Berkeley. Paratypes are de-
posited in the American Museum of Natural History, New York; Califor-
nia Academy of Sciences; California Insect Survey; Canadian National
Collection, Ottawa; Los Angeles County Museum; and U.S. National
Museum, Washington, D.C.

Coptodisea powellella Opler, new species

Male. Length of forewing 1.5 to 1.7 mm (reared).

HoLotyPE MALE. Length of forewing 1.6 mm. Forewing: Basal half metallic
silver; distal half with pattern as follows: a basal orange patch extending from costa
almost to inner margin; followed distally by two opposing triangular patches of silver,
one on costal margin, one on inner margin, apices not quite joining; these patches
narrowly margined with black basally and distally; costal area orange distal to tri-
angular patches except for subapical black margined silver patch and black apex.
Dorsal area black distal to silver patch on inner margin. Abdomen: Blackish dor-
sally, silver ventrally. Genitalia: As in Figs. 9, 10, 12, 14, 15 (drawings from para-
type, PAO prep. 323); uncus shallowly bilobed at tip.

Female. Length of forewing 1.6 to 1.9 mm (reared).

ALLOTYPE FEMALE. Length of forewing 1.7 mm. External features as described for
male. Genitalia: As in Figs. 28, 29, 30. Comparison with genitalia of other oak-
feeding Coptodisca was not made. Thus, the included drawings serve only to portray
the structures which may later prove to be of diagnostic value.

Type Material. Holotype male: California, Descanso Ranger Station, San Diego
County, 31 March 1961, reared from Quercus agrifolia Neé, J. Powell lot 61C22,
emerged 21 April 1961, collected by J. Powell. Allotype female: same data except
emerged 12 April 1961. Paratypes: 662 4, 839 2 as follows: California.—Contra
Costa Co.: Clayton, 6 mi. SE, 19, II-2-68, r.f. QO. agrifolia, J. Powell lot 68B18,

VoLUME 25, NUMBER 3 195

Figs. 1-4. 1, Lithocolletis antiochella Opler, ¢, right forewing, holotype; 2,
Coptodisca powellella Opler, ¢, right forewing, Descanso Ranger Station, San Diego
Co., Calif. 31 March 1961 r.f. Quercus agrifolia JAP 61C22 emgd. 18 April 1961,
J. Powell collector; 3, Cameraria wislizeniella Opler, ¢, right forewing, holotype;
4, Coptodisca querciella Braun, ¢, right forewing, paratype, Cornville, Yavapai Co.,
Ariz. rf. Quercus emoryi B1241 emgd. 1 May 1925, H. Brisley collector.

emgd. III-20-68 (P. Opler); Cowell, 19, II-28-68, rf. QO. agrifolia, J. Powell lot
68B193, emgd. IV-10-68 (P. Opler). Los Angeles Co.: Santa Monica Mts., 3é 6,
3Q 9, III-17-69, rf. QO. agrifolia, J. Powell lot 69C68, emgd. IV-3/10-69 (P. Opler).
Monterey Co.: Arroyo Seco Camp, 4¢ 6, II-12-68, rf. QO. agrifolia, J. Powell lot
68B53, emed. III-27/1V-5-68 (P. Opler). Placer Co.: Penryn, 14, II-13-68, rf. OQ.
wislizenii, J. Powell lot 68C14, emgd. IV-8-68 (P. Opler). Riverside Co.: Mountain
Center, 1¢, III-16-68, r.£. Q. wislizenii var. frutescens, J. Powell lot 68B127, emegd.
III-17-68 (P. Opler). San Bernardino Co.: Forest Home, 1 mi. W, 3¢ ¢, 52 Q, III-
29-68, rf. O. wislizenii var. frutescens, J. Powell lot 68C77, emgd. IV-27/V-7-68
(P. Opler). San Diego Co.: Descanso Regr. Sta., 56 6, 69 9, III-31-61, II-15-68,
r.f. QO. agrifolia, J. Powell lots 61C22, 68B110, emgd. IV-10/21-61, II-14/23-68 (P.
Opler, J. Powell). Warner Springs, 1 mi. NW, 32¢ 6, 42929, II-15-68, rf. Q.
agrifolia var. oxyadenia, J. Powell lot 68B118, emgd. III-12/IV-5-68 (P. Opler).
Santa Barbara Co.: Los Prietos, 1 mi. E, 106 ¢, 129 9, II-13-68, r.f. Q. agrifolia,
J. Powell lot 68B76, emgd. III-14/IV-5-68 (P. Opler); Santa Cruz Island, Prisoner’s
Harbor, 46 6, 59 9, III-14-69, rf. QO. agrifolia, J. Powell lot 69C24, emgd. IV-8/
14-69 (P. Opler, J. Powell); Santa Cruz Island, U. C. Field Sta., 1¢ 6, 792 9, III-
15-69, r.f. QO. agrifolia, J. Powell lot 69C48, emgd. III-27/IV-6-69 (P. Opler).

196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 5-8. 5, Acrocercops insulariella Opler, 9, Central Valley, Santa Cruz Is-
land, Santa Barbara Co., Calif., 28 April 1966 r.f. Quercus agrifolia JAP 66D45,
emgd. 31 May 1966, J. Powell and J. Wolf collectors; 6, Neurobathra bohartiella
Opler, 6, Strawberry Canyon, Alameda Co., Calif. 30 July 1968, r.f. Quercus agrifolia
JAP 68G73, emgd. 13 August 1968, P. Opler collector; 7, Caloptilia agrifoliella Opler,
2, two miles southest of Inverness, Marin Co., Calif., 13 April 1968, rf. Quercus
agrifolia JAP 68D1333, emgd. 12 May 1968, P. Opler collector; 8, Lithocolletis
sandraella Opler, 6, Descanso Ranger Station, San Diego Co., Calif., 31 March
1961, r.f£. Quercus agrifolia JAP 61C24, emgd. 9 April 1961, J. Powell collector.

The only previously described western oak-feeding member of this
genus is C. querciella Braun. In her description of C. querciella, Braun
(1927) delineates only the maculation features of the adult and states
that the host is an evergreen species of Quercus. Through Dr. Braun’s
courtesy I was able to examine leaf mines as well as a male paratype from
Cornville, Yavapai County, Arizona, the type locality. The leaf which
harbored the mines of C. querciella proved to be Quercus emoryi, a
species which is consubgeneric with the hosts of C. powellella.

C. powellella differs from C. querciella in features of forewing macula-
tion. The former has extensive orange markings and restricted black
markings, while the latter has less extensive orange and more extensive
black ( Figs. 2, 4).

The most obvious differences in the male genitalia are in the aedeagi
and apparent aedeagal sheaths of the two species. The “aedeagal sheath”
of C. powellella is proportionately larger than that of C. querciella and

VOLUME 25, NUMBER 3 197

13

Figs. 9-13. Male genitalia of Coptodisca. 9, C. powellella Opler, dorsal aspect,
aedeagus removed; 10, C. powellella, left valva, ventral aspect; 11, C. querciella
Braun, left valva, ventral aspect; 12, C. powellella, aedeagus and “aedeagal sheath”
lateral aspect; 13, C. querciella, aedeagus and “aedeagal sheath”, lateral aspect.

198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

is of a somewhat different shape, while other apparent differences are
shown by the tips of the aedeagi (Figs. 12, 13).

Acrocercops insulariella Opler, new species

Male. Length of forewing, 3.7 to 4.5 mm (reared).

HouoryPE MALE. Length of forewing 4.2 mm. Head: Maxillary palpus with
outer side black scaled, inner side white scaled. Labial palpus primarily white
scaled basally, black scaled distally. Antenna longer than wing, uniformly blackish
gray. Vestiture of front mixed brownish-gray and gray white. Vestiture of vertex
brownish-gray. Thorax: Scaling of dorsum and tegulae brownish-gray; scaling of
pleural and ventral areas predominantly white with some light gray scales inter-
mixed. Prothoracic leg with femur dirty white, tibia gray, tarsomeres gray black
ringed with white; mesothoracic leg with femur white ventrally, blackish-gray dor-
sally, tibia black with white basoventrally and a white ring around middle, tarsomeres
black, ringed with white; metathoracic leg with femur white, tibia white ventrally,
black dorsally, tarsomeres black, variously marked with white. Forewing: Ground
mixed brown and orangish brown; several small patches of five to ten white scales as
follows: seven along costal margin, four along inner margin. Underside dark brown.
Fringe brown with two patches of white scales at distal end of costa, a few black-
tipped white scales at wing tip, and a few white scales in fringe at distal end of
inner margin. Hind wing: Upperside, underside, and fringe gray brown. Abdomen:
Gray black dorsally, white ventrally. Genitalia: As in Figs. 17, 18 (drawings from
Paratype, PAO prep. no. 139); valvae with dorsal margin straight, margin of cucullus
slightly convex distally to meet costal margin at bluntly subtriangulate tip, inner face
densely covered with setae on distal .8; tegumen .75 length of valvae, distal margin
quadrate in ventral view, weakly setate; saccus short; aedeagus with one pointed
cornutus at tip.

Female. Length of forewing 4.3 to 4.7 mm (field collected).

ALLOTYPE FEMALE. Length of forewing 4.5 mm. External features as described
for male. Genitalia: Aphophyses anteriores slightly recurved. Ductus bursae with
rib-like sclerotizations at .66 from opening. Signum a caliper-shaped sclerotization.

Type Material. Holotype male: California, Santa Cruz Island, Central Valley,
Santa Barbara County, 28 April 1966, reared from Quercus agrifolia Neé, J. Powell
lot 66D45, emerged 31 May 1966, collected by J. Powell, A. Slater, and J. Wolf.
Allotype female, California, Santa Cruz Island, Eagle Canyon east of Prisoner’s Har-
bor, 9 June 1966, collected by J. A. Powell. Paratypes: 66 6, 329 as follows:
California.—Santa Barbara Co.: Santa Cruz Island, Prisoner's Harbor Creek, 4é ¢,
19, IV-28-66, rf. QO. agrifolia, J. Powell Lot 66D45, emgd. V-26/31-66 (J. Powell,
A. Slater, J. Wolf); Santa Cruz Island, Eagle Cyn., E of Prisoner's Harbor, 2é 6,
12, VI-9-68 (J. Powell).

Acrocercops insulariella differs from A. affinis Braun (1918), the only
previously described oak-feeding species which occurs in the west, by
features of forewing maculation and the male genitalia. Through the
courtesy of Dr. Braun I was able to examine male and female paratypes
from Camp Baldy, San Bernardino County, California, the type locality.

Although the two species are closely related there are slight but con-
stant differences. The forewing of A. affinis has a pale white patch on
the inner margin two-fifths from the base which has quadrate darkened

patches to either side. These features are not present in individuals of A.
insulariella.

VOLUME 25, NUMBER 3 199

{
1
1
1
‘ii
|
‘|
ol

23 240 ee

Figs. 14-24. Male genitalia. 14, Coptodisca powellella Opler, tip of uncus, dor-
sal aspect; 15, C. powellella, transtilla, dorsal aspect; 16, Coptodisca querciella
Braun, transtilla, dorsal aspect; 17, Acrocercops insulariella Opler, ventral aspect,
aedeagus removed; 18, A. insulariella, aedeagus, lateral aspect; 19, Acrocercops affinis
Braun, ventral aspect, aedeagus removed; 20, A. affinis, aedeagus, lateral aspect; 21,
Neurobathra bohartiella Opler, ventral aspect aedeagus removed; 22, N. bohartiella,
aedeagus, lateral aspect; 23, Caloptilia agrifoliella Opler, ventral aspect, aedeagus re-

moved; 24, C. agrifoliella, aedeagus, lateral aspect.

200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

The male genitalia of the two species differ in that the uncus of A.
insulariella is truncate with two prominent long setae distally, while the
uncus of A. affinis is subacuminate distally and has several short setae.

Neurobathra bohartiella Opler, new species

Male. Length of forewing 4.5 to 4.9 mm (reared).

Ho.LoryPE MALE. Length of forewing 4.7 mm. Thorax: Scaling of dorsum and
tegulae tan; scaling of pleural and ventral areas silvery gray. Prothoracic leg with
femur blackish, tibia silvery gray, tarsomeres white basally, black distally; meso-
thoracic leg with femur white anteriorly, blackish posteriorly, tarsomeres white
ventrally, black dorsally with black distally; metathoracic leg with basal .25 of femur
blackish, distal .75 white, tibia white ventrally, gray dorsally, tarsomeres black
basally, white distally. Forewing: Ground gray brown; ground scaling on distal
third of wing tipped with tan; a black mark on costa .3 from base, three narrow,
black, outwardly directed streaks on costa near .75 from base; three narrow, white,
outwardly directed streaks on costa at from .5 to .8 from base; a white, subapical
patch on costa; a large white streak curving outwardly from inner margin .75 from
base. Underside brown. Inner fringe at wing tip black; outer fringe at wing of black-
tipped white scales giving fringe a black-white-black pattern; remainder of fringe
brown. Abdomen: Brownish-gray dorsally, white ventrally. Genitalia: As in Figs.
21, 22 (drawings from Holotype, PAO prep. no. 173); valvae almost half as broad
as long, truncated distally; two short, stout, triangulate projections at distal end, the
ventral largest; costal margin of valva straight.

Female. Length of forewing 4.1 to 4.7 mm (reared).

ALLOTYPE FEMALE. Length of forewing 4.3 mm. External features as described
for male. Genitalia: Anterior margin of seventh segment a heavily sclerotized ring.
Anterior portion of corpus bursae sclerotized with small finger-like projections in-
teriorly.

Type Material. Holotype male: California, Strawberry Canyon, Alameda County,
30 July 1968, reared from Quercus agrifolia Neé, J. Powell lot 68G73, emerged 13
August 1968, PAO prep. no. 173, collected by P. Opler. Allotype female: California,
Golden Gate Park, San Francisco County, 26 February 1968, reared from OQ. agrifolia,
J. Powell lot 68B181, emerged 6 March 1968, collected by P. Opler. Paratypes:
468, 499 as follows: California.—Alameda Co.: Berkeley, 16, III-1-59 (J.
Powell); Strawberry Cyn., 1¢, VII-30-68, r.f. QO. agrifolia, J. Powell lot 68G73,
emgd. VIII-13-68 (P. Opler). Los Angeles Co.: Westwood Hills, 19, IV-41, Acc.
309 (R. M. Bohart). Mendocino Co.: Yorkville, 6 mi. SE, 19, V-18-66 (A. J.
10-69, r.f. Q. agrifolia, J. Powell lots 68B33, 68LB, 69B6, emed. II-19-68, XII-15-
Slater). San Francisco Co.: Golden Gate Park, 2¢ 4, 19, II-6-68, XI-15-68, II-10-
69, rf. Q. agrifolia, J. Powell lots 68B33, 68L5, 69B6, emed. II-19-68, XII-15-68,
II-7-69 (P. Opler). Santa Barbara Co.: Refugio Beach, 3 mi. N, 19, VII-4-65 (J.

Powell).

Neurobathra bohartiella is apparently the only representative of the
genus in California. Neurobathra strigifinitella (Clemens), the only
other described species in the genus, occurs in eastern North America.
The genitalia of N. bohartiella differ from that of N. strigifinitella as
figured by Vari (1961) as follows: the costal margin of the valva of N.
bohartiella is straight while that on N. strigifinitella is slightly concave.
Additionally the upper tooth on the distal margin of the valva is relatively
smaller for N. bohartiella than for N. strigifinitella.

VOLUME 25, NUMBER 3 201

The life cycles of the two species differ in that pupae of N. strigifinitella
undergo an obligatory pupal diapause (fide T. N. Freeman), while the
life cycle of N. bohartiella lacks any form of diapause.

Caloptilia agrifoliella Opler, new species

Male. Length of forewing 6.2 to 7.0 mm (reared ).

HoLotyPe MALE. Length of forewing 6.8 mm. Head: Maxillary palpus yellow
scaled inwardly, mixed yellow and russett outwardly. Labial palpus yellow scaled
basally, mixed yellow and russet distally. Antenna slightly longer than forewing;
scape orange scaled; flagellar segments yellowish basally, blackish distally. Vestiture
of front and vertex brownish orange. Thorax: Scaling of dorsum and tegulae brown-
ish orange with bluish iridescence in reflected light; scaling of pro- and meso-
thoracic pleural areas orange; scaling of metathoracic pleural areas yellowish. Pro-
thoracic leg with femur yellowish-orange; tibia and basal tarsomeres orangish-brown;
remainder of tarsomeres silvery white, narrowly ringed with black distally. Meso-
thoracic leg with femur and tibia dark orangish-brown; tarsomeres white, narrowly
ringed with black distally. Metathoracic leg with femur white-scaled, tibia white
ventrally, gray dorsally; tarsomeres white, ringed narrowly with dark gray distally.
Forewing: Brownish-orange with ill-defined yellowish orange areas on costa; a
bluish sheen in reflected light. Underside gray black narrowly edged with orange.
Fringe gray black. Hind wing: Upperside, underside, and fringe gray black. Ab-
domen: Scaled gray black dorsally, white ventrally. Genitalia: As in Figs. 23, 24
(drawings from Paratype, PAO prep. no. 148); valva becoming broader distally,
slightly truncated at tip, twice as broad distally as at base; dorsal margin flaring;
ventral margin with shallow convex lobe at .5 from base, inner face of lobes with
about 15 peg-like setae arranged in three rows; anterior corner of valva with a patch
of about 30 stout, pointed setae; distal half of valva covered with long hair-like
setae on inner face. Tegumen sclerotized, broader and longer than vinculum, tri-
angulate in distal half, basal half with ventrally directed flaps. Tuba analis mem-
branous, projecting beyond vinculum, with weakly sclerotized median ridge. Saccus
as long as length of valva, narrow, acuminate distally. Aedeagus tubular, pointed at
tip, lacking cornuti. Coremata of two groups of hair-like scales, one group with
scales more than 3 X length of the other.

Female. Length of forewing 6.2 to 6.8 mm (reared).

ALLOTYPE FEMALE. Length of forewing 6.6 mm. Externally as described for
male. Genitalia: Ninth segment ventrally with posteriorally directed peg-like projec-
tion. Eighth segment with a deep membranous invasion just posterior and ventral
to apophyses anteriores. Signa two slightly curved, cone-like projections.

Type Material. Holotype male: California, one mile southeast Inverness, Marin
County, 2 May 1969, reared from Quercus agrifolia Neé, J. Powell lot 69E76, emerged
19 May 1969, collected by P. Opler. Allotype female: California, two miles south-
east Inverness, Marin County, 13 April 1968, reared from Q. agrifolia, J. Powell lot
68D133, emerged 12 May 1968, collected by P. Opler. Paratypes: 54 ¢, 109 9 as
follows: California.—Alameda Co.: Strawberry Cyn., 1¢, 22 9, IV-11-68, rf. Q.
agrifolia, J. Powell lot 68D124, emgd. IV-7/V-25-68 (P. Opler). Marin Co.: In-
verness, 1 mi. SE, 16, 59 9, V-10-68, V-2-69, r.f. Quercus agrifolia, J. Powell
lots 68E19, 69E76, emgd. V-30-68, V-19/23/69 (P. Opler, J. Powell); Inverness, 2
mi. SE, 32 @, IV-13-68, r.f. QO. agrifolia, J. Powell lot 68D133, emgd. V-12/14/68
(P. Opler). Monterey Co.: Bixby Cyn., 16, VII-27-48 (J. W. Tilden). San
Francisco Co.: Golden Gate Park, 1¢, VII-30-67, r.f. Q. agrifolia, J. Powell lot
67G22, emgd. VIII-25-67 (P. Opler). Santa Barbara Co.: Santa Cruz Island, Pri-
soner’s Harbor, 1 6, VI-7-66 (R. L. Langston).

202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Caloptilia agrifolia can be distinguished from Caloptilia reticulata
(Braun), the only other described oak-feeding species of the genus oc-
curring in California, on the basis of life cycle features, larval shelter,
maculation, and genitalia. C. agrifoliella is normally single-brooded, but
is facultatively double-brooded, feeds only on young foliage, and has a
pupal diapause which usually lasts through the summer, fall and winter.
The larval shelter is a rolled leaf. C. reticulata is homodynamically mul-
tiple-brooded, feeds on older foliage, and lacks any type of diapause. The
larval shelter consists of a singly folded leaf. The vestiture on the dorsum
of the thorax and tegulae of C. agrifoliella is unicolorous; and the fore-
wing is variously colored orangish, tan, or rust colored with faint light
areas and black marks. The male genitalia is distinguished by the long
acuminate saccus, rounded valvae, and triangulate tegumen. The vesti-
ture on the dorsum of the thorax of C. reticulata is reddish-orange, while
that of the tegulae is straw-colored, and the forewing is reddish-orange
with numerous straw-colored splotches. The male genitalia are dis-
tinguished by the short saccus, truncated valvae, and rounded tegumen.

The names Gracilaria sanguinella Beutenmiller, Gracilaria nigristrigella
Beutenmiiller, Gracilaria ruptistrigella Beutenmiller, Gracilaria shastella
Beutenmiiller described in 1888, and Gracilaria fuscoochrella Beuten-
miller described in 1889 were synonymized by Ely (1917) under Gra-
cilaria alnivorella Chambers. McDunnough (1946) restricted the name
alnivorella to an alder feeding species of eastern North America and did
not include Beutenmiiller’s names in the synonymy of that species.

Remnants of the Beutenmuller holotypes are located in the U.S. Na-
tional Museum and were examined by J. A. Powell. The present state
of these types is as follows:

G. sanguinella—*1358 California.” No abdomen. No host given.

G. nigristrigella—*1333 California.” All that remains of the type is one
hind wing. Collected on oak tree.

G. ruptistrigella—*‘1327 California.” The specimen is broken in half.
No host given.

G. shastella—No abdomen. No host given.

G. fuscoochrella—No abdomen. No host given.

These names could apply to any of a number of Californian Caloptilia
species which feed on Acer, Alnus, Lithocarpus, or Quercus. Because
features of the genitalia and host plant are the main criteria for assigning
specific status to the moths of this genus and because the above listed
Beutenmuller names cannot be evaluated on either basis, I treat them
as nomena dubia.

VOLUME 25, NUMBER 3 203

A = ‘

ae
AONE he
Ss ~ : ea Aor
\ re Xe /
= a —s Raa é
ECR ff
LaF
} i
b d
F _
fo
f 4
= E /
*: ~ i i
a * \ q
mek, oh i
\ : j rh
. : Ni
~
: \ de ZG:
Las a
; aS:
os >»
ys
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Figs. 25-27. Male genitalia, ventral aspect. 25, Lithocolletis antiochella Opler;
26, Lithocolletis sandraella Opler; 27, Cameraria wislizeniella Opler.

204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Lithocolletis antiochella Opler, new species

Male. Length of forewing 2.7 to 3.9 mm (reared).

HoLotTyPE MALE. Length of forewing 3.5 mm. Head: Maxillary palpus brownish-
tan scaled, antenna tan, slightly shorter than length of forewing costa. Vestiture of
front shining white; vestiture of vertex, brownish-tan. Thorax: Scaling of dorsal
portion and anterior half of tegulae brown, whitish at anterior margin; posterior
halves of tegulae white-scaled. Pleural and ventral areas with shining white scaling.
Legs with shining white scaling except for black scaling on: dorsal halves of femora,
tibia, and tarsi of pro- and meso-thoracic legs. A tuft of long white scales on ventral
half of tibiae. Forewing: Brown, shining gold in reflected light; a white stripe from
base through median portion of wing along basal .4 of length, a white dorsal streak
at .2 distance from base, directed outwardly and joining basal median stripe; two
pairs of opposite white streaks narrowly edged with black inwardly, the basal pair .33
from base directed sharply apically, not quite joining, the dorsal streak largest, distal
pair small, triangulate, each extending .33 width of wing; a median streak of black
scales on termen not quite reaching wing tip; wing tip edged narrowly with black.
Underside brown. Fringe pale tan. Hind wing: Upperside pale tan. Underside
brownish-tan. Fringe pale tan. Abdomen: Tan dorsally, whitish ventrally. Geni-
talia: As in Fig. 25 (photograph from Paratype, PAO prep. no. 246); valvae asym-
metrical, both valvae with width gradually increasing distally, dorsal margins strongly
decurved; right valva ending with a short, stout spine directed posteroventrally,
ventral margin straight; left valva with ventral margin with a concavity just basal
to distal end; saccus short, triangular, well sclerotized; transtilla well sclerotized;
aedeagus straight, narrow, and sclerotized basally, broadened and weakly sclerotized
for distal .33; uncus acutely triangular; flap of ninth sternite long, subacute at tip.

Female. Length of forewing 2.7 to 3.9 mm (reared).

ALLOTYPE FEMALE. Length of forewing 3.2 mm. Externally as described for male.
Genitalia: Eighth segment completely membranous. Posterior margin of seventh seg-
ment concave ventrally at opening of ductus bursae. Signum bursae a small round
sclerotization with stout seta-like projection.

Type Material. Holotype male: California, 2 miles east of Antioch, Contra Costa
County, 28 February 1968, reared from Quercus agrifolia Neé J. Powell lot 68B198,
emerged 5 March 1968, collected by P. Opler. Allotype female: same data. Para-
types: 358 6, 5122 as follows: California—Alameda Co.: Strawberry Cyn.,
146, VII-30-68, r.f. Q. agrifolia, J. Powell lot 68G71, emgd. II-12-69 (P. Opler).
Contra Costa Co.: Antioch, 2 mi. E, 246 6, 229 9, II-2-68, II-28-68, r.f. QO. agri-
folia, J. Powell lots 68B2, 68B198, emgd. II-15/19-68, III-1/10-68 (P. Opler), 74 4,
27 ¢ 6, II-28-68, III-26-69, OQ. agrifolia assoc. (P. Opler, J. Powell, C. Slobodchikoff );
Cowell, 2¢ 6, 19, II-28-68, II-16-69, r.f. O. agrifolia, J. Powell lots 68B192, 69B16,
emgd. III-7-68, II-21-69 (P. Opler); Russell Farm, 4 mi. NE Orinda, 19, III-6-70,
Q. agrifolia assoc. (P. Opler). Los Angeles Co.: Westwood Hills, no abdomen, IV-
41, rf. Q. agrifolia (R. M. Bohart). Ventura Co.: Newbury Park, 14, IX-5-69,
r.f. QO. agrifolia, J. Powell lot 69J28, emgd. XII-15-69 (P. Opler).

Its life history, maculation, and genitalic features set Lithocolletis
antiochella apart from all other members of the genus. Among the species
treated by Braun (1908) in her revision of the North American Litho-
colletis, Lithocolletis diaphanella Frey and Boll appears to be most closely
related in external appearance to L. antiochella. L. antiochella differs
from L. diaphanella in that the vestiture of the palpi, antennae, and ver-
tex are brown, not white; that there are three dorsal white streaks in-
stead of two; and that the forewings are brown in color as opposed to

VOLUME 25, NUMBER 3 205

peas

i 1

' U !

[ ae 1
raw !

1 !

} ' ;

J ‘
ro
' ’ |
sie

( a4 I

/
6 1
On 1
16 \ i

Wess

\\f%o if 00

\ a foe \
=U \

df

Figs. 28-30. Coptodisca powellella Opler, female genitalia. 28, Genital appara-
tus, ventral aspect; 29, posterior portion of eighth segment, ventral aspect; 30, tip of
abdomen showing distal portion of ovipository apparatus.

golden brown. A comparison of the genitalia of these species was not
carried out.

The male genitalia of L. antiochella have a well sclerotized diaphragma,
asymmetrical valvae which are broader distally than basally. The eighth
abdominal segment of the female is completely membranous.

The larvae feed on young foliage in the spring and then undergo a
pupal diapause of about ten months.

Lithocolletis sandraella Opler, new species

Male. Length of forewing 2.9 to 4.7 mm (reared).

HoLotyPE MALE. Length of forewing 3.8 mm. Head: Maxillary palpus white
scaled. Antenna tan, slightly longer than .5 costal length. Vestiture of front shining
white; vestiture of vertex light orange-tan with a few scattered white scales latero-
dorsally. Thorax: Pronotal scaling and tegulae orange-tan with two narrow white

206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs, 31-35, Female genitalia. 31, Acrocercops insulariella Opler, latero-ventral
aspect; 32-33, Neurobathra bohartiella Opler, ventral aspect; 34, Caloptilia agri-
foliella Opler, lateral aspect, corpus bursa removed; 35, C. agrifoliella corpus bursa.

VoLUME 25, NUMBER 3 207

streaks along junction of inner margin of tegulae with dorsum of thorax. Pleural
and ventral areas with white scaling. Legs white-scaled except dorsal halves of pro-
femur and protibia fuscous scaled. Forewing: Ground orange-tan, a thin white
stripe from base through median portion of wing for .35 length, a white mark at
middle of costal margin directed postero-apically and terminating at point on median
of width and .66 length from base, a patch of black-tipped scales running from inner
termination of costal white mark to subapical area; at apex fringe with basal row
of black-tipped scales. Underside dark tan. Fringe tan with long tufi of black scales
at apex. Hind wing: Upperside white. Underside tan. Fringe white. Abdomen:
Black dorsally, white ventrally. Genital segments with tan scaling externally. Geni-
talia: As in Fig. 26 (photograph from Paratype, PAO prep. no. 250); valvae sym-
metrical, broad at base (width about .33 length), narrowing distally with tip bluntly
acuminate, a crescentic hollow area on ventral margin near apex containing patch
of short, stout setae, a stout filament-like seta .25 length of valva at inner edge of
hollow area directed ventrodistally; flap of ninth sternite long, parallel-sided, grad-
ually rounded apically.

Female. Length of forewing 3.5 to 5.5 mm (reared).

ALLOTYPE FEMALE. Length of forewing 4.3 mm. External features as described
for male except dorsum of abdomen pale tan-scaled. Genitalia: As in Fig. 37.
Apophyses posteriores with small triangulate projections .33 from base; apophyses
anteriores short. Genital plate enlarged, heavily sclerotized, inserted into seventh
segment. Signum bursa an elongate lip-like sclerotization.

Type Material. Holotype male: California, Descanso Ranger Station, San Diego
County, 31 March 1961, reared from Quercus agrifolia Neé, J. Powell lot 61C24,
emerged 5 April 1961, collected by J. Powell. Allotype female: same data. Para-
types: 1366, 2022 as follows: Alameda Co.: Berkeley, 1¢, III-7-69, r.f. QO.
agrifolia, J. Powell lot 69C5, emgd. III-20-69 (P. Opler); Berkeley Hills, above Old
Tunnel, 19, VIII-25-68, r.f. QO. agrifolia, J. Powell lot 68H16, emgd. IX-3-68 (P.
Opler). Contra Costa Co.: Cowell, 1¢, II-28-68, rf. Q. agrifolia, J. Powell lot
68B194, emgd. IV-5-68 (P. Opler); Tilden Park, 1¢, IV-16-69, rf. QO. agrifolia,
J. Powell lot 69D31, emgd. IV-27-69 (P. Opler). Kem Co.: Keene, 29 2, X-1-67,
r.f. QO. wislinzenii, J. Powell lot 67K9, emgd. X-7/20-67 (P. Opler). Los Angeles Co.:
Malibu Cyn., 5 mi. N. Malibu, 19, III-17-69, rf. QO. agrifolia, J. Powell lot 69C59,
emegd. III-2-69 (P. Opler); Topanga Cyn., 1¢, II-17-69 r.f. Q. agrifolia, J. Powell
lot 69C58, emgd. III-31-69 (P. Opler). Monterey Co.: Carmel, 19, X-2-27 (L. S.
Slevin); Jolon, 17.5 mi. W, 3é¢ 6, 19, III-19-69, r.f. QO. wislizenii, J. Powell lot
69C87, emgd. IV-6-69 (P. Opler). Orange Co.: Silverado, 1 ¢, IIJ-30-68 (P. Opler).
San Benito Co.: Limekiln Cyn., SW Paicines, 19, IV-23-69, rf. Q. agrifolia, J.
Powell lot 69D55, emgd. V-7-69 (P. Opler). San Diego Co.: Descanso Rgr. Sta.,
266, 29 9, I-31-61, rf. O. agrifolia, J. Powell lot 61C24, IV-7/14-61 (J. Powell),
19, III-31-61 (R. L. Langston); Guatay, 1 mi. W, 29 2, IX-13-68, r.f. Q. agrifolia
var. oxyadenia, J. Powell lot 68J73, emgd. IX-21/24-68 (P. Opler); Honey Spgs. Rd.,
16, X-4-67, rf. QO. agrifolia, J. Powell lot 67K51, emgd. X-31-67 (P. Opler). San
Francisco Co.: Golden Gate Park, 19, IV-5-68, r.f. Q. agrifolia, J. Powell lot 68D49,
emgd. IV-23-68 (P. Opler). San Luis Obispo Co.: La Panza Camp, 19, IV-25-68,
r.f. QO. agrifolia, J. Powell lot 68D131, emgd. V-7-68 (P. Opler). Santa Barbara Co.:
Santa Cruz Island, Prisoner’s Harbor, 2? 2, III-14-69, r.f. Q. agrifolia, J. Powell lot
69C23, emgd. III-31-69 (P. Opler). Santa Clara Co.: Herbert Crk., 3 mi. W New
Almaden, 192, IV-25-69, rf. O. agrifolia, J. Powell lot 69D71.1, emgd. V-7-69 (P.
Opler). Ventura Co.: Newbury Park, 26 6, 32 @, IX-5-69, rf. Q. agrifolia, J.
Powell lot 69J27, emgd. IX-6/11-69 (P. Opler).

Lithocolletis sandraella is quite unlike any other North American spe-
cies of the genus. Perhaps Lithocolletis inusitatella Braun is the species

208 JouRNAL OF THE LEPIDOPTERISTS SOCIETY

=<

aot
saat sea =

38

Figs. 36-38. Female genitalia, ventral aspect. 36, Lithocolletis antiochella Opler;
37, Lithocolletis sandraella Opler; 38, Cameraria wislizeniella Opler.

VOLUME 25, NUMBER 3 209

with which L. sandraella should be compared. Both L. sandraella and L.
inusitatella have Quercus agrifolia and Q. wislizenii as their only hosts.
L. sandraella lacks the metallic bronze sheen and apical black spot dis-
played by the forewings of L. inusitatella. L. sandraella differs strikingly
from L. inusitatella in genitalic features as well. The valvae of L. sand-
raella are symmetrical while those of L. inusitatella are asymmetrical.
Additionally, L. sandraella is bivoltine, while L. inusitatella is univoltine.

Cameraria wislizeniella Opler, new species

Male. Length of forewing 3.1 to 4.7 mm (reared).

HOLOTYPE MALE. Length of forewing 3.8 mm. Head: Maxillary palpus white
scaled. Antenna as long as costal length; scape white ventraily, brownish orange dor-
sally, flagellum fuscous scaled dorsally, tan ventrally. Vestiture of front shining
white; vertex white mesally, narrowly margined with brownish orange laterally.
Thorax: Scaling of dorsum broadly white mesally, brownish-orange laterally; tegulae
brownish-orange; pleural and ventral areas with shining white scaling. Prothoracic
leg black dorsally, white ventrally, tarsomeres ringed with white; mesothoracic leg
white, tarsomeres black, ringed with white; metathoracic leg with femur white, tibia
white ventrally, brownish-orange dorsally, tibia and tarsa predominantly white, with
small black mark on dorsal surface at junction of tibia and tarsus and junction of first
and second tarsomeres. Forewing: Ground bronzy brownish-orange; an outwardly
directed narrow white streak on costa at .5 forewing length from base, narrowly edged
at outer margin with black, a broken narrow white stripe extending from tornus to
wing tip edged anteriorally with white. Underside blackish fuscous. Fringe brown-
ish-orange apically, gray white posteriorally. Hind wing: Upperside shining gray
white. Underside gray. Fringe gray white. Abdomen: Gray black dorsally and
laterally, white ventrally. Genitalia: As in Fig. 27 (photograph from Paratype, PAO
prep. no. 231); valvae symmetrical, narrow, broadest basally, sinuous, falcate apically;
a single row of setae on inner face of valva extending from base to .5 length; a
patch of longer setae on inner face of valva at ventral margin .66 from base, uncus
long, acuminate, two slender setae at distal tip; flap of ninth sternite long, distal
half triangulate, deeply cleft distally to separate two truncate lobations.

Female. Length of forewing 2.4 to 4.3 mm (reared).

ALLOTYPE FEMALE. Length of forewing 3.4 mm. As described for male except:
inner margin of forewing narrowly edged with white on basal half (this character-
istic present on many males). Dorsal scaling of abdomen pale tan. Genitalia: As in
Fig. 38 (drawing from Paratype, PAO prep. no. 338); opening of ductus bursae sur-
rounded by posterad bilobed projection. Signum bursae consisting of a small tri-
angular sclerotization.

Type Material. Holotype male: California, Knight's Ferry, Stanislaus County,
20 March 1968, reared from Quercus wislizenii A. D. C., J. Powell lot 68C46, emerged
24 March 1968, collected by J. T. Doyen. Allotype female: California, Woodacre,
Marin County, 15 March 1968, reared from Quercus wislizenii var. frutescens En-
gelm., J. Powell lot 68C23.1, emerged 8 April 1968, collected by P. Opler. Paratypes:
376 6, 2622 as follows: California.—Contra Costa Co.: Antioch, 2 mi. E,
306 6, 219 9, II-2-68, II-28-68, IV-7-68, V-26-68, r.f. Q. agrifolia, J. Powell lots
68B4, 68B199, 68D56, 68E47, emgd. II-14/19, III-7/26, IV-13/25, VI-6/14 (P.
Opler); Clayton, 4 mi. SE, 192, V-26-68, rf. Q. wislizenii, J. Powell lot 68E65,
emgd. by II-11-69 (P. Opler); Clayton, 8 mi. SE, 1¢, II-2-68, rf. Q. wislizenii, J.
Powell lot 68B13, emgd. II-20-68 (P. Opler); Mt. Diablo, summit, 3849’, 19,
IV-13-69, rf. QO. wislizenii var. frutescens, J. Powell lot 69D141, emgd. IV-24-69

210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(P. Opler). Marin Co.: Woodacre, 36 6, III-15-68, r.f. Q. wislizenii var. frutescens,
J. Powell lot 68C23.1, emgd. IV-5/8-68 (P. Opler). Monterey Co.: Jolon, 17.5 mi.
W, 14, III-19-69, r.f. O. wislizenii, J. Powell lot 69C86, emgd. III-31-69 (P. Opler).
Placer Co.: Penryn, 16, 29 9, IJI-13-68, r.f. Q. wislizenii, J. Powell lot 68C15,
emed. III-23/IV-8-68 (P. Opler). San Bernardino Co.: Forest Home, 1 mi. W, 1°,
III-29-68, r.f. O. wislizenii var. frutescens, J. Powell lot 68C79, emgd. IV-23/V-1-68
(P. Opler). Stanislaus Co.: Knight’s Ferry, 1 ¢, IlI-29-68, r.f. QO. wislizenii, J. Powell
lot 68C46, emgd. IV-10-68 (J. T. Doyen).

Among described species of the genus, the closest relative of Cameraria
wislizeniella is Cameraria mediodorsella (Braun). These moths are very
similar in maculation and are difficult to separate on that basis, but the
two have distinctive male genitalia. The male genitalia of C. wislizeniella
differs by the distally expanded valvae, the relatively uncurved aedeagus,
and the attenuate flap of the ninth sternite.

The life cycle of C. wislizeniella is typitied by the absence of diapause
while that of C. mediodorsella has a pupal diapause over the winter
months.

Acknowledgments

I wish to thank the following for the loan of material from their private
or institutional collections: P. H. Armaud, Jr., California Academy of Sci-
ences, San Francisco; A. F. Braun, Cincinnati, Ohio; D. R. Davis, U.S.
National Museum, Washington, D.C.; J. A. Powell, California Insect
Survey, Berkeley; R. O. Schuster, University of California, Davis; J. W.
Tilden, San Jose, California.

J. A. Powell provided helpful suggestions and reviewed the manuscript
of this paper. Miss Susan McDonald made the illustrations of adult
Lithocolletis antiochella and Cameraria wislizeniella. My wife Sandra
provide secretarial assistance.

Funds supporting field and laboratory work during this study were
provided by National Science Foundation grants GB4014 and GB6813X
under the direction of J. A. Powell.

Literature Cited

BEUTENMULLER, W. 1888. On North American Tineidae. Ent. Amer. 4: 29-30.

1889. On the North American Tineidae. Ent. Amer. 5: 9-10.

Braun, A. F. 1908. Revision of the North American species of the genus Litho-
colletis Hiibner. Trans. Amer. Ent. Soc. 34: 269~357.

——— 1918. New species of Microlepidoptera. Can. Ent. 50: 229-236.

wae eee Descriptions of new Microlepidoptera. Trans. Amer. Ent. Soc. 53:
J1I—LYY,

ny, C. R. 1917. A revision of the North American Gracilariidae from the stand-
point of venation. Proc. Ent. Soc. Wash. 19: 29-77.

KuMATA, T. 1963. Taxonomic studies on the Lithocolletinae of Japan. (Lepidop-
tera: Gracillariidae). Insecta Matsumurana 25: 1-90, 26: 1-88.

VoLUME 25, NUMBER 3 2

Kuroxo, H. 1961. The genus Antispila from Japan, with descriptions of seven new
species ( Lepidoptera, Heliozelidae). Esakia 3: 11-24.

McDunnoucn, J. H. 1946. Gracillariid studies (Gracillariidae, Lepidoptera).
Can. Ent. 78: 91-95.

Vari, L. 1961. South African Lepidoptera. Vol. 1. Lithocolletidae. Transvaal
Museum. Mem. 12.

OBITUARY

SS

ANTONIE JOHANNES THEODORUS JANSE
1877-1970

On June 12th, 1970, at the age of ninety-three, Dr. A. J. T. Janse, a great Lepidop-
terist and a man of remarkable learning and ability, passed away. For almost seventy
years he collected, reared, studied, and described Lepidoptera, a privilege given to
very few. All by himself, he built up a unique collection of the Heterocera of southern

Africa.

912, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Dr. Janse was born in The Hague, the Netherlands, where he received his education.
He emigrated to South Africa in 1899, and became a teacher in Waterval (Nuwe
Smitsdorp). He started his entomological work under the most primitive conditions,
but his boundless enthusiasm enabled him to overcome these difficulties and his pains-
taking ceaseless collecting laid the foundation for all his far-reaching scientific work
of later years. In the early days he did his collecting on foot, later by donkey cart.
His was pioneer work, for at that time there was no extensive collection of Lepidoptera
in South Africa.

In 1905 he accepted the post of lecturer at the Transvaal Normal College, and this
marked the turning-point in his early career, since it enabled him to take up ento-
mology in earnest and to become a scientist. The facilities available to him in Pretoria
made it possible for him to undertake systematic research.

In 1921-1922 he visited Europe and worked at the museums in London, Leiden
and Berlin, comparing and identifying many hundreds of specimens from his collection.
In recognition of his work he was presented in London with the Joicey collection of
Pyralidae, which he brought back to Pretoria.

After his return, Janse was appointed honorary professor of Systematic Entomology
at the Transvaal University College. He held this post until his retirement in 1937.
In 1925 the University of South Africa awarded him the honoris causa degree of
Doctor of Science.

The most rewarding acknowledgment of Janse’s entomological achievements came
in 1945, when the South African Government bought his collection, library, and equip-
ment, and placed them in the care of the Transvaal Museum, he himself being ap-
pointed Honorary Curator of Heterocera. However, because the museum had no space
to house Janse’s collection, it was decided to leave it at his house, where his laboratory
was also located, and to transfer the museum’s collection of Heterocera there instead,
thus uniting the two collections. The Government’s action was of vital importance
for the future of Janse’s life work, which was now secure; the museum provided the
necessary funds to buy urgently needed drawers and other equipment. An assistant
was appointed to continue his work eventually. In 1953 space became available
for the collection at the museum, but Janse continued to work in his laboratory, using
material on loan from the main collection.

During the greater part of his life Janse had to do all his entomological work during
holidays and in his spare time. Nevertheless, he found time to publish a substantial
series of papers, and in 1932 began on his outstanding series “The Moths of South
Africa.” But it was not until his retirement a few years later that he was able to give
his full attention to the publication of this valuable series. Up to the time of his death
he had published over forty papers and eight large volumes of the Moths series. He
described more than five hundred new species, for which he also prepared detailed
drawings of external structures and genitalia of both sexes. He always maintained that
such drawings should be made a compulsory part of any description of new taxa.

Before his death, Janse had completed the manuscript for the next volume of “The
Moths of South Africa” and had almost finished another volume on the types of South
African Microlepidoptera described by the late E. Meyrick.

During his association with the Transvaal Museum, Janse made three extensive col-
lecting trips by motor car and trailer; although he was already over seventy-five, he
travelled with his wife as his only assistant. He continued his research with unflagging
zeal through the years, until at last a serious illness put an end to his activities about
a year before his death.

Janse’s frequent letters were always striking and written in a firm hand; a year ago
he wrote “so long my hand and eyes are still good, can the work go on.” These letters
breathed such vitality, enthusiasm, and trust in his work that they conveyed something
fresh, simple, and deeply encouraging, especially when one remembered the writer’s

age
Ue.

Only a few are blessed with such a wealth of enthusiasm for their work. Few can

VOLUME 25, NUMBER 3 2A3

devote themselves to it so wholeheartedly. Add to this unusual stamina and insight,
and one has the stuff of which Janse’s great generation of Lepidopterists was made. His
must have been an enviable, full, and happy life.

L. VAri, Transvaal Museum, Pretoria, South Africa and A. Diaxonorr, Leiden Mu-
seum, The Netherlands.

PRELIMINARY NOTES ON THE BUTTERFLIES OF ROOSEVELT COUNTY,
| NEW MEXICO

Roosevelt County is situated on the central eastern border of New Mexico, in the
southern portions of the Great Plains. Its primary industry is agriculture, but despite
this, it has one of the most interesting butterfly faunas of the state.

I have been able to make only four trips to this area, two in 1965 and two in 1966.
As a result of these four trips, I have recorded 38 species, plus two more seen. One
other is recorded from the county; Megathymus coloradensis elidaensis Stallings,
Turner, and Stallings, the type locality of which is located southwest of Elida. Of
the thirty-eight species of which I have specimens, four have not been taken by me
from anywhere else in the state. Of the remaining thirty-four, twelve are represented
by four records or less in the state. One of the species seen but not taken was a bat-
tered specimen of Papilio cresphontes Cramer, a species which has also never been
recorded elsewhere in the state to my knowledge.

The main locality in which I have collected is located six miles west of Portales,
along New Mexico Highway 88. This is an extremely unattractive place for the Lepi-
dopterist, being situated at the junction of four farm fields, two to the north of the
highway and two to the south. On the south side of the road, there is a short row of
elm trees. The tallest of these is about 25 feet high. It was on one of these elms that
a larva of Polygonia interrogationis (Fabricius) was collected. The larva unfortunately
died later, but there is little doubt as to its identity. One adult was also seen, but es-
caped capture. The only other record the author has of this species in New Mexico
is at his home in Albuquerque.

The majority of the butterflies were collected at roadside flowers, which are es-
pecially numerous in the fall. The area between the edge of the fields and the road
is quite wide (at least 75 feet), so there is a large patch of them.

Following is a list of the thirty-eight species collected at this locality on September
12, 1965; September 18-19, 1965; June 2-3, 1966, and June 11, 1966. Known state
records are marked with an (*) and unusual records are marked with an (!).

Hesperiidae

*1. Lerodea eufala (Edwards); Sept. 19, (2 males).

12. Amblyscirtes eos (Edwards); June 3, (1 male).

3. Atalopedes campestris (Boisduval); Sept. 19, (1 male, 1 female); June 2, (1
female ).

4. Hesperia uncas uncas (Edwards); Sept. 19, (1 female); June 2, (11); June 3,
(17 )s Iria JUL, (Le

15. Hylephila phyleus (Drury); June 2, (1).

6. Pholisora catullus (Fabricius); Sept. 12, (1); Sept. 19, (1); June 3, (2); June
i, Cae

7. Pyrgus communis (Grote); Sept. 12, (1); Sept. 18, (1); Jume 2, (1); June 3,
(4)= June 11) (1).

137.

38.

. Vanessa virginiensis (Drury); June 11, (

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Papilionidae

Battus philenor philenor (Linnaeus); June 3, (1); June 11, (1).
Papilio polyxenes asterius Stoll; June 3, (5); June 11, (2).

Pieridae

Pieris protodice Boisduval & Le Conte; Sept. 19, (2 males); June 3, (2).
Colias eurytheme Boisduval; Sept. 12, (1 male, 1 female); Sept. 19, (2 males,
5 females, 3 female alba); June 3, (1); June 11, (6).

Phoebis sennae eubule (Linnaeus); June 11, (1 male).

Kricogonia lyside (Godart); June 3, (1 male, 5 females).

Eurema mexicana (Boisduval); Sept. 19, (1 male).

Eurema lisa Boisduval & Le Conte; Sept. 12, (2); Sept. 19, (3 females).
Eurema nicippe (Cramer); Sept. 19, (2); June 3, (6); June 11, (1).

. Nathalis iole Boisduval; Sept. 12, (2); Sept. 19, (6); June 2, (1); June 3, (1);

June lea (Gle)e

Lycaenidae

Strymon melinus franki Field; Sept. 19, (2); June 2, (7); June 3, (8); June
(Gye

. Brephidium exilis (Boisduval); Sept. 19, (1).

Leptotes marina ( Reakirt); June 2, (2); June 3, (3).
Hemiargus isola alce (Edwards); Sept. 12, (2); Sept. 19, (2); June 2, (5);
une B, (15).

. Plebejus acmon lutzi dos Passos; June 3, (2).

Libytheidae

Libytheana bachmanii (Kirtland); June 3, (1).
It is not known whether this specimen is larvata (Strecker) or typical bachmanii.
I have been unable to locate the specimen.

Nymphalidae

Anaea andria Scudder; Sept. 12, (1); Sept. 18, (3).

Mestra amymone ( Menetries ); Sept. 12, (1).

Four other specimens were seen on this date, but unfortunately I was sans net.
See Masters, (1970).

}. Vanessa atalanta (Linnaeus); Sept. 19, (1); June 11, (4).
)

Vanessa cardui (Linnaeus); Sept. 18, (1); June 3, (3); June 11, (2).

. Nymphalis antiopa (Linnaeus); Sept. 18, (1); Sept. 19, (1).
. Chlosyne lacinia adjutrix Scudder; Sept. 12, (5); Sept. 18, (1); Sept. 19, (11);

June 3, (1).

. Phyciodes texana texana (Edwards); June 2, (1)

Phyciodes phaon (Edwards); Sept. 19, (1 male, 1 female).

Phyciodes picta canace Edwards; Sept. 12, (1); Sept. 19, (1); June 2, (2);
June 3, (4).

Phyciodes vesta (Edwards); June 3, (1).

Texola elada ulrica (Edwards); June 3, (1).

Sabon claudia (Cramer); Sept. 19, (1); June 2, (1); June 3, (2); June
eels):

Agraulis vanillae incarnata (Riley); June 3, (1); June 11, (2).

Danaidae

Danaus gilippus strigosus (Bates); June 3, (4).

MicHAr FE, Toriver, 1612 Indiana NE, Albuquerque, New Mexico.

VoLUME 25, NuMBER 3 AM,

OBITUARY

:

SN

JOHN ADAMS COMSTOCK
Jan. 30, 1883-Dec. 26, 1970

Entomologists will learn with regret of the death, on December 26, 1970, of Dr.
John Adams Comstock, at his home in Del Mar, California. Past President and Hon-
orary Life Member of the Lepidopterists’ Society, he would have been 88 years old
One janes.) 1971.

His interest in Lepidoptera started at a very early age and continued throughout
his long and very productive life. He published over 220 papers in various journals,
and he wrote and published The Butterflies of California.

A complete biography and bibliography will be published in the 25th anniversary
publication of the Lepidopterists’ Society in 1972.

Lioyp M. Martin, 19 Meadow Lane, Route 1, Prescott, Arizona.

216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

FOREST-PRAIRIE TRANSITIONS AND THE ADMIXTURE
OF BUTTERFLY FAUNAS

Kurt JOHNSON!

Terms referring to general types of ecological communities, such as
forest, prairie, tundra, or montane communities, are used by ecologists to
describe or categorize the character of environments. Likewise, they are
used in description of types of species inhabiting such areas. Terms
such as prairie butterfly, or forest butterfly, though sometimes vague in
meaning, are common descriptive terms in Lepidopterology.

Transitions from prairie communities to forest communities differ in
degree of abruptness from place to place. The escarpment affords a wide
area of interspersement of these types of communities, while an abruptly
rising mountain range may leave only a small amount of margin between
forest and grassland. It seems obvious that the degree of interspersement
of forest and prairie has a counterpart in the extent of forest, prairie, or
forest-prairie types of species making up an area's faunal admixture.

Recent studies of butterfly faunas in escarpments along the western
edges of the Great Plains, where forested areas are interspersed with
prairie (Johnson and Nixon, 1967; Johnson, 1971) suggest extensive
sympatry of butterfly species from various type of ecological communities.
Hence, an investigation into the effect of forest-prairie interspersement
on faunal admixtures seemed invited.

I therefore created a scheme by which types of species indicating the
forest, prairie, or forest-prairie ecology might be designated and _ their
percentage composition in various faunas analyzed.

Materials and Methods

It was deemed important to conduct the study in areas representing
intergradations between two extremes. Hence, I selected the Bighorn
Mountains of Wyoming (an area where transition from grassland to forest
is abrupt ), the scantily forested escarpments of the Long Pine Recreation
Area in north-central Nebraska, and several areas between them. The
latter included the Laramie Mountains of Wyoming, the Front Range of
Colorado, the Black Hills of South Dakota, the Pine Ridge of Nebraska,
and the escarpments along the Niobrara River cuttage in Nebraska. The
location of these study areas is presented in Figure 1. Since nearly all of
these areas have had recent specific faunal studies, a list of species for
each area was conveniently obtained within the criteria cited concerning

Present address: Novitiate, Order of the Holy Cross, West Park, New York.

VoLUME 25, NUMBER 3 217

Se.

SOUTH DAKOTA
WYOMING

NEBRASKA

COLORADO

Fig. 1. Locations of the seven study areas. 1, Bighorn Mountains; 2, Black Hills;
3, Laramie Mountains; 4, Front Range; 5, Pine Ridge; 6, Niobrara River cuttage; 7,
Long Pine Recreation Area.

them. Because the faunas of these separate areas are not unknown to lepi-
dopterists and because the lists were compiled from a variety of sources,
it was not deemed important to the study to include them, except to note
that the species were organized according to dos Passos (1964). The
number of species in each fauna is listed below each column on the den-
drogram. The sources of the faunal lists are summarized below.

The faunal list of the Pine Ridge in Nebraska came from Johnson and
Nixon (1967). That of the Niobrara River cuttage and Long Pine Recre-
ation Area from Johnson's The Butterflies of Nebraska (1971). The Wy-
oming lists were obtained from the literature (DeFoliart, 1956; Nabokov,
1953) and the extensive personal research of Richard Hardesty (Douglas,
Wyoming) and John S. Nordin (Webster, South Dakota). Hardesty and
Nordin also assisted in compiling the faunal list of the Black Hills of South
Dakota. The Colorado list was culled from county records in Brown
(1957).

Since it is difficult to construct an exact criterion to designate which

218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

species are found “within” any physiographic area, a consistent decision
was made to define mountain areas as all parts which rose above the low-
lands, that is, having a higher elevation than the surrounding plains. Simi-
larly, regarding river cuttages, this criterion included all areas below the
surrounding plains, and in escarpment situations, that transition between
surrounding lowlands and table lands above. Although criteria of this
type cannot be completely exact, the amount of variation within the defi-
nitions seem not too dangerous, since it leaves the amount of general in-
terspersement of forest and prairie dependent on the abruptness of the
transition between dominance of forest or prairie. In this study these
criteria grouped the study areas into three general categories: escarp-
ments, with complete interspersement (Pine Ridge, Niobrara cuttage,
Long Pine Rec. Ar.), areas of moderate transition (Black Hills, Front
Range, Laramie Mts.) and an area of abrupt transition (Bighorn Mts.).
That the abruptness of transition affects consistently the types of species
present is the important thesis of this investigation.

The second problem was developing definitions for forest butterfly,
prairie butterfly, and forest-prairie butterfly. It was most important that
the technique be clear and as unbiased as possible. To satisfy this, Mr.
Hardesty and I separately gave each species one of three labels: forest,
prairie, or forest-prairie. The criteria used were based upon questions:
“Must one go to the forest (or prairie, or either) to find the species?” and
if this could not be definitive, “In what environ does the foodplant grow?”
The success of this method is evident since Hardesty and I disagreed on
the designation in only a few cases. In these I made the redesignation on
the grounds of further consideration of the original criteria. With lists
of all the resident Papilionoidea defined into these three categories for
each study area, the percentage of each category in each fauna was com-
puted. Results are summarized in Figure 2, which presents dendrograms
of the percentage composition of each fauna.

To facilitate the accuracy of sampling, the Hesperiidae were omitted
since they are poorly known in some of the collection areas. It should be
noted, however, that this omission tends to depress the percentage of
prairie species in the admixture since many of the skippers are of that
type. Also omitted were any tundra forms inhabiting a study area. The
use of general categories like forest or prairie as defined in this study
seems to allow for this. To give the Colorado list a realistic “Front Range”
integrity, the list from that area was limited to Larimer and Boulder Coun-
ties. Also, nowhere were the faunal resemblances between the faunas
involved (these are calculable after Long, 1963)
forest (deciduous or coniferous) designated.

nor was the type of

y

VOLUME 25, NUMBER 3 219

17 FP
75 50 : 594 PSs 46, F
“g 3 14 15 19 p

70 64 98 107 81 43 24

Fig. 2. Graphic representation of faunal admixture in each study area. Forest-
prairie taxa (FP) illustrated above, clear; Forest taxa (F) represented in middle,
stipled; Prairie taxa (P) represented below, clear. 1, Bighorn Mountains; 2, Black
Hills; 3, Laramie Mountains; 4, Front Range; 5, Pine Ridge; 6, Niobrara River cuttage;
7, Long Pine Recreation Area.

Figures below each bar represent number of species in each fauna.

Results

The results are summarized in Figure 2. Each bar of the graph is
arranged to illustrate change in number of prairie taxa (below, clear),
forest-prairie taxa (above, clear), and forest taxa (middle bar, stipled).

The order of the graphs corresponds with the degree of transition eco-
logically proceeding from the most abrupt (Bighorm Mts., 1) to the most
gradual (the Long Pine Rec. Ar., 7). Note that both the Front Range
sample and Laramie Mountains sample are represented though they re-
flect no real difference in abruptness. This serves to illustrate similarity
in types of admixture in two very similar ecological situations.

The portions of each graph representing percentages of species indi-
cating certain types of ecological situations show a consistent pattem
relative to the abruptness of the ecological transition from prairie to forest.
The general correlations are as follows:

920 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Increase in abruptness, corresponding with less interspersement of
prairie and forest, results in:

1. increase in forest indicative species
2. decrease in prairie indicative species
3. decrease in forest-prairie indicative species

Decrease in abruptness, corresponding with more interspersement of
prairie and forest, has the opposite result in all cases.

Consistent with the general ecological concept that ecotones often
harbor a major portion of the species of biota in an area, results indicate
that the complete interspersement of forest and prairie biomes in the
escarpments promotes a dominance of those areas by forest-prairie types
of species (see Long Pine Rec. Ar., 7). In fact, with decrease in abruptness
of transition the amount of forest-prairie species increases at a greater rate
than purely prairie taxa, indicating the pressure of the marginal ecology
on the faunal admixture. Discussion of the relative roles of prairie and
forest in the margin is very difficult, but it appears that the factor of forest
may dominate an area. This is illustrated by the fact that even a semblance
of woodland, such as the scattered stands of trees in the Long Pine area
affords large numbers of forest species a habitat, probably by its affording
the general flora of the area a divergence. This conforms to the obser-
vation that food-plant diversity in the forest and especially in the ecotonal
areas is usually greater than on the surrounding mixed-prairie. The role
of the Pine Ridge forests in influencing the admixture of the Nebraska
butterfly fauna (Johnson, 1971) is an example of this on a large scale. The
relationship of types of species and the habitats they utilize invites new
thinking into problems of taxonomic and distributional evolution.

Conclusions

The study illustrates that when types of species indicating certain eco-
logical conditions are designated realistically, their percent admixture in
a fauna varies consistently with degrees of spatial transition in an areas
ecology.

Though the general assumption is simple, the analysis of it seems justi-
fied since the assumption is usually made without any thought to the
parameters or criteria for its having meaning.

Hence, an effort was made to give definition to categories that are
usually general or non-definitive. The paper points out that such a defin-
itive study can be made, and hopefully offers criteria on which similar
questions can be investigated.

VOLUME 25, NUMBER 3 yp )|

Acknowledgments

I am grateful to Dr. C. A. Long (Wisconsin State University, Stevens
Point) for his suggestions concerning the project, and to Mr. Richard
Hardesty (Douglas, Wyoming) and Dr. John S. Nordin (Webster, South
Dakota ) for their aid in compiling the faunal lists.

Literature Cited

Brown, F. M., D. Err anp B. Rotcer. 1957. Colorado butterflies, Proc. Denver
Mus. Nat. Hist., Denver.

DeFourart, G. R. 1956. An annotated list of southeastern Wyoming Rhopalocera.
Jour. Lepid. Soc. 10: 91-101.

pos Passos, C. F. 1964. A synonymic list of the Nearactic Rhopalocera. Lepid.
Soc. Mem. 1.

Jounson, K. anp E. S. Nixon. 1967. The Rhopalocera of northwestern Nebraska.
Amer. Mid. Nat. 78: 508-528.

Jounson, K. 1971. The butterflies of Nebraska. Lepid. Foundation (in press).

Lone, C. A. 1963. Mathematical formulae expressing faunal resemblance. Trans.
Kansas Acad. Sci. 66: 138-140.

Nasoxoy, V. 1953. Butterfly collecting in Wyoming, 1952. Jour. Lepid. Soc. 7:
49-52.

BOOK REVIEW

Morus anp How To REAR THEM, by Paul Villiard. 1969, 242 pp. + i-xiii, profusely
illustrated with half tones. Funk and Wagnalls, 380 Madison Avenue, New York.
Price $10.00 U.S.

This is a volume that everyone interested in rearing the larger moths, particularly
the sphingids and saturniids, should have on his reference shelf. One hundred and
seventy-seven species are discussed under the headings of family [name], popular
name, range, availability, preferred foodplant, diapause [stage], ova, larvae, rear-
ing requirements, cocoon and adult. The adult, the mature larva, the cocoon or
pupa, and usually the egg of each of the native and exotic species discussed are
illustrated in half-tone reproduction.

Although an introductory section is devoted to a detailed description of the
spreading procedure, some of the adults illustrated have not been spread with any
great skill and many of these appear to have become wet and matted at some stage.
The author-photographer seems to have a penchant for dark backgrounds in his
illustrations and in the case of dark specimens or those with translucent wings results
are not pleasing. The usage of generic names is often not current but this causes no
difficulty.

The book is a highly enjoyable one and I thoroughly recommend it to all interested
in the natural histories of the larger moths.

D. F. Harpwicx, Editor.

929, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

UNUSUAL- BUTTERFLIES IN NORTHERN PENNSYLVANIA

Following a few years’ collecting as a boy in Tioga County, Pennsylvania in the mid
1920s, I again began to take an amateur interest in butterflies about ten years ago.

The rewakening of active interest really began when I discovered a colony of larvae
of Battus philenor on a Dutchmans Pipe Vine (Aristolochia macrophylla) at my home
in Wellsboro in late July 1959. This common southern butterfly is rare in northern
Pennsylvania so the larvae were cared for until the adults emerged in late August. No
other wild larvae or adults have been seen here since.

On May 15, 1960, I caught a single Pieris virginiensis while I was on a hiking trip
in an open mixed hardwood forest about 8 miles southeast of Wellsboro. This led to
an active search and since that time a number of colonies of varying sizes have been
located in the area.

On July 6, 1963, while exploring an acid bog for orchids and other plants ( Polygonia
ophioglossoides, Calopogon pulchellus, Sarracenia purpurea, Drosea rotundifolia, Meny-
anthes frifoliata and Vaccinum macrocarpon are present) about thirty miles southwest
of Wellsboro, I noted a small blue-gray butterfly which proved to be Lycaena epixanthe.
In a mountain-top meadow near this bog, Arthur Shapiro scored a first for the area by
netting a male Colias interior on July 11, 1968.

On May 19, 1968, while I was walking along a grassy woods road, about eight miles
southeast of Wellsboro, in an area now grown up to mixed hardwood brush, a fresh
female Erora laeta literally dropped into the road in front of me. Later that same year,
on July 29th, a worn male was collected and another sighted on blossoming hardhack
(Spirea tomentosa) in a nearby wet field.

Last season (1970) on June 25th, July Ist and July 5th, in two locations about
twenty miles apart north and west of Wellsboro respectively, I collected a total of
ten Thymelicus lineola, all males. These insects were feeding on the blossoms of
common milkweed (Asclepias syriaca) but were very wary and difficult to approach.
No females have been collected.

A number of large colonies of Chlosyne harrisii liggetti Avinoff occur here, flying
from early June to early July.

The Tioga-Potter-Lycoming County area in north-central Pennsylvania can be clas-
sified as Transitional Zone with tendencies toward Canadian Zone in the higher eleva-
tions. Mountains are rounded with maximum elevations of approximately 2200’. Valley
bottoms are about 1300’ elevation. It is mostly heavily forested with second-growth
hardwoods, principally maples, beech, cherry, ash, basswood and some oaks. There
are scattered hemlocks and some red and white pine.

The presence of the L. expixanthe acid bog can possibly be attributed to pre-glacial
times and the peculiar geological history of the area. Geologists agree that the edge
of the last glacier was nearby and that previous to that time the region drained north-
ward into the St. Lawrence River. The glacier halted this flow, however, and an
entire new drainage system developed. A new water passage was cut through the then
existing mountain barrier to the south forming what is now known as “Pennsylvania's
Grand Canyon,” a steep-sided gorge about fifty miles long and one thousand feet deep,
draining into the Susquehanna River and thence into the Atlantic Ocean at Chesapeake
Bay. In Potter County, about forty miles west of Wellsboro, there is now a three-way
watershed, with drainage dividing into flowages to the north (St. Lawrence River), to
the southwest (Mississippi River), and to the southeast (Susquehanna River).

GrorcE F. Patrerson, Wellsboro, Penn.

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NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
for material in excess of 20 printed pages, at the rate of $17.50 per page. Address
all correspondence relating to the Journal to: Dr. D. F. Hardwick, K. W. Neatby
Bldg., Central Experimental Farm, Carling Ave., Ottawa, Canada.

Contributors should prepare manuscripts according to the following instructions;
failure to do so will result in unnecessary delay prior to publication.

Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
margins, on one side only of white, 84% x 11 inch paper. Authors should keep a
carbon copy of the MS. Titles should be explicit and descriptive of the article’s
content, including the family name of the subject, but must be kept as short as
possible. The first mention of a plant or animal in the text should include the
full scientific name, with authors of zoological names. Underline only where italics
are intended in the text (never in headings). References to footnotes should be
numbered consecutively, and the footnotes should be typed on a separate sheet.

Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
heading LireRAturRE Crrep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: All photographs and drawings should be mounted on stiff, white
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Volume 25 1971 Number 4

NOTES ON GREGARIOUS ROOSTING IN TROPICAL
BUTTERFLIES OF THE GENUS MORPHO

ALLEN M. Younc!

Organization for Tropical Studies, Inc.,
Apartado 16, Universidad de Costa Rica, Costa Rica, C. A.

Recent literature on general theories in community ecology has stressed
the evolution of highly specialized patterns of behavior in species that in-
habit tropical rain forests, (Margalef 1968; Miller 1969; Slobodkin and
Sanders 1969). It is held that the stable (e.g., Levins 1968) or predictable
nature (e.g., Levins and MacArthur 1967; Slobodkin and Sanders, 1969 )
of the physical environment in the tropics allows for organisms to cue in on
the subtle features of the habitat, and adjust very well by developing very
specialized and highly evolved patterns of behavior related to survivorship
in low-density populations.

Clearly, studies of behavioral patterns associated with reproduction,
survivorship, and daily activity patterns in natural populations of tropical
organisms are needed to assess the generality of such theoretical consid-
erations. This present paper summarizes observations on repeated gre-
garious nocturnal roosting in the tropical butterflies, Morpho amathonte
centralis and Morpho granadensis polybaptus (Lepidoptera: Morphinae ).
The observed consistency and duration of this behavior in males of these
large butterflies indicates that it may have selective value with respect
to (1) efficient exploitation of food sources and (2) reservation of the
bulk of daily energy budgets for courtship activities. With the recent ex-
ception of a discovery of this phenomenon in a supposedly tasteful nymph-
aline, Marpesia bernia, (Emmel and Benson 1971), gregarious nocturnal
roosting habits were known mainly in distasteful species of tropical butter-
flies (e.g., Poulton 1930; Carpenter 1933; Crane 1955; 1957; Owen and
Chanter 1969) with the adaptive value of such behavior being that the
noxious or distasteful qualities of the butterflies en masse is a much more
effective predator-deterring behavior than if individuals roosted singly

1 Present address: Department of Biology, Lawrence University, Appleton, Wisconsin 54911.

294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

(Crane 1955). The study of Emmel and Benson (1971) and the present
paper suggests that different species of tropical butterflies, both distasteful
and tasteful forms, have evolved such behavior as different adaptive
strategies. Unlike the interpretation of such behavior in other butterflies
(e.g., Emmel and Benson 1971), it is unlikely that gregarious roosting in
Morpho butterflies acts as a mechanism against predatory attacks.

Observations

Four roosts of adult male Morpho amathonte were discovered during
January and early February 1970 in a lowland tropical rain forest (“La
Selva” ) located in northeastern Costa Rica. These discoveries were made
in conjunction with observations on the feeding and flying activities of
these butterflies, and these studies are summarized elsewhere (Young
1971a). All of the butterflies seen at each roost turned out to be those
that had been marked previously at their daily feeding sites (natural ac-
cumulations of fallen fruits of the canopy tree, Coumarouna (= Diptyerx )
oleifera ( Papilionaceae—“Almendro” ); the group of butterflies associated
with a feeding site were the same individuals that roosted together during
the night. There was no exchange of individuals among the widely-
separated four feeding sites studied and observations were made to deter-
mine the extent of roost faithfulness, in addition to survivorship and re-
cruitment rates. At the time of discovery of these roosts, they consisted of
5, 2, 2 and 5 individuals (with the order being roosts 1, 2, 3 and 4 to cor-
respond with feeding sites 1, 2, 3 and 4 discussed in Young 1971a).

One roost of adult male Morpho granadensis was discovered during
May 1970 in the montane tropical rain forest located near Cuesta Angel
in central Costa Rica. This roost initially consisted of 14 individuals and
over a 10-day period, all of these butterflies were marked in order to
determine roost faithfulness, survivorship, and recruitment rates. Unlike
Morpho amathonte, these individuals were not marked at their feeding
sites since these were not known. However, at La Selva, a few individuals
of this species were seen roosting with Morpho amathonte and these
butterflies were also observed. Marking was facilitated in this species as
they were easily netted after 5 pm (CST). In both species, an enamel base
paint, “F'lo-Paque,” was used to mark butterflies; each butterfly was given
2 distinct markings on the undersides of the wings: one mark to denote
roosting site, and another mark to denote individual number.

The four roosts of Morpho amathonte were observed for five consecutive
months, with observations being made on several days per month. The
roost of Morpho granadensis was observed for 3.5 months and not as fre-
quently as those of Morpho amathonte. At the time of marking, the wing
condition of individuals (tattering, loss of scales, fading) was noted in

VoLUME 25, NuMBER 4

iw)
i)
Ol

order to study changes in the age-structure of the adult population; sex
was also determined. The latter was easily done since males are much
brighter and possess different markings than females, and thus sexes could
be distinguished at a distance.

In the case of Morpho amathonte, all four roosts were situated in low,
open, secondary-growth vegetation encompassed by closed-canopy forest.
These roosting sites were very close to feeding sites and typically no more
than 35 meters away. Adults usually roosted on the upper surfaces of
large-sized leaves of various plant species, and in vertical positions. Wings
were always kept tightly closed while roosting, and against the background
of dark green foliage and shadows, the butterflies were difficult to see
even at close distances. Adults were widely spaced with the nearest dis-
tance between two individuals being a few feet. It was never seen that
more than one adult rested on a leaf; perhaps in this respect roosts of
Morpho butterflies are structurally different than those of other species.
Emmel and Benson (1971) report on the occurrence of several individuals
of Marpesia bernia roosting together on the undersides of a leaf. Un-
doubtedly size of the insect is an important factor in determining the
relative proximity of individuals in a roost; Morpho butterflies are very
large (average wingspan 25.5 cm in both species) and perhaps require
more space on a per individual basis. Furthermore, adult males are ag-
gressive along their flight paths (Young 197la) and this may be an im-
portant factor in determining the distances among roosting individuals.
Aggressive interactions among males at roosting sites have not been
observed.

The roost of adult male Morpho granadensis was located on branches of
trees that were overhanging a river (Rio Sarapiqui) and individuals
hanged from the upper surfaces of leaves. Whereas the roosts of Morpho
amathonte took on a more circular configuration due to the shape of the
openings in the forest used, the roost of Morpho granadensis was oblong
in shape in that it extended along the river bank and included various
species of trees. However, individuals were again widely-separated. In-
dividuals roosted about 1.5 meters above the water and in the case of
Morpho amathonte, individuals roosted from 2 to 5 meters above the
ground. In two of the roosts of Morpho amathonte, individuals of Morpho
granadensis were seen over long periods of time. At one roost (roost 2)
there were 3 individuals of Morpho granadensis, and at a second roost
(roost 4) there were 2 individuals. Like the Cuesta Angel roost, all of
these individuals were males.

A summary of the temporal patterns of roost occupancy are given in
Table 1. As seen for adult aggregations at feeding sites (Young 1971]a),
there was virtually no turnover in adult males of Morpho amathonte at

926 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TaBLE 1. Evidence for high roost faithfulness* in adult Morpho butterflies in Costa
Rican tropical rain forests.

Number? of Butterflies
Months NP Locality Roost No. Females Males Total

Morpho amathonte

Feb. 14 La Selva 1 0 5 5
Mar. 20 0 5 5
Apr. 18 0 6 6
May 22 0 6 6
June* 3) 0 5 5
Feb. 14 La Selva 2 0 2 2
Mar. 20 0 DD 2)
Apr. 15 0 3 3
May 20 0 3 3
June® 6 0 2 YD
Feb. 14 La Selva 3) 0 2 2,
Mar. 18 0 2 Y
Apr. 7 0 2 2
May 19 0 2 2
June* 5 0 2; 2
Feb. 13 La Selva 4 0 5 5
Mar. 20 0 5) 5
Apr. 18 0 6 6
May 20 0 6 6
June* 5 0 5 5
Morpho granadensis
Apr. 8 Cuesta Angel ] 0 14 14
May iLL 2 16° 18
June iL 0 10 10
July° 8 I ise 14

* Defined here as the continuous occupancy of the same roosting site(s) over a long period of
time (several months).

>» N is the number of days of observation at roots each month.

© These are means; standard errors were very small and are not included in the table.

4 For Morpho amathonte, there was only one week of observation during early June; for Morpho
granadensis, there was only 8 days of observation during early July.

® Increments in adults numbers was due to appearances of newly-emerged males at the roost.

their nocturnal roosts. Turnover was more prevalent with Morpho grana-
densis (Table 1).

The time of departure and arrival of the butterflies at the roosts each
day was highly variable and depended upon local weather conditions.
This was particularly true for Morpho granadensis. On overcast after-
noons, individuals arrived usually between 4-6 pm, whereas on sunny
afternoons, arrival was usually restricted to 5:30 to 6:15 pm. It is of in-
terest to describe the pattern of arrival of individuals at the Cuesta Angel
roost of Morpho granadensis. The roost was located along a river bank at

4

VoLUME 25, NUMBER 4 pay

the base of a steep gorge covered with closed-canopy tropical rain forest.
Standing on the opposite of the river and directly in front of the roost site,
with the aid of binoculars, the pattern of arrival could easily be observed
for these very large and conspicuous butterflies. On all days of observa-
tion (a total of 54), the arrival pattern was the same and fascinating: one
by one, and following each other at very close intervals, the butterflies
would lazily fly down to the roost from the upper reaches of the gorge.
They formed an imaginary line of blue objects as they flew down the
gorge over the tops of the trees. They always approached the roost from
one side and several meters away; then they would fly along the river bank
until they reached the roost. On a sunny day, all individuals were in the
roost within 25 minutes; the time involved was longer on overcast days but
the pattern was the same. By being stationed very close to the roost, it
was possible to observe the actual order (of marked individuals) of ar-
rivals; this pattern was highly variable each day. Departure occurred
usually between 8:30-9:30 am each day and individuals flew off in dif-
ferent directions. In Morpho amathonte, arrival was staggered and indi-
viduals flew in from different directions. Departure was of a similar pat-
tern and the roosts were usually vacated by 7:00 am each day. No con-
sistent order of arrival and departure for the marked individuals of this
species could be determined. Young (197la) suggests that adult males
of this species elect to roost overnight in open, exposed areas in the forest
so that they may receive the early sun ray to assist their bodies becoming
sufficiently warmed-up for flight. This in turn, is related to the diurnal
activity pattern of this species: adult males feed from 7-8:30 aM, fly on
consistent flight paths from about 8:30 am to 11:30 aM, and either partake
in courtship activity along flight paths beyond this hour or rest near
feeding sites before moving to nocturnal roosts between 5:00 and 6:00 pm
(Young 197la). Under such a diurnal activity pattern, it is expected that
arrivals at roosts would be inconsistent, depending upon whether in-
dividuals were involved in courtship or resting near feeding sites.

On the other hand, both adult male and females of Morpho granadensis
concentrate the bulk of their feeding activity to the late afternoon hours
(3:00-6:00 pm) and presumably engage in courtship activity earlier in
the day (Young 1971b). Like Morpho amathonte, adults of Morpho
granadensis aggregate at feeding sites (at La Selva, these can be the
same ones used by Morpho amathonte) (Young 1971b). The cessation
of feeding with advancing nightfall or undesirable weather conditions may
account for the “group return” pattern of arrivals to roosts in this species;
since all individuals are involved in the same activity and at the same time,
it is expected that they would be more consistent in arriving at roosts.

While it is clear that there is no movement of individuals once they are

228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

settled in roosts, if a roost is approached (by human observer ) to within
about 2 meters of a resting individual, that individual will fly away for a
short distance. When the disturbance is stopped, the butterfly will return
to its original perch usually within 15 minutes. Depending on the direction
of flight upon disturbance, an individual can either cause other resting
individuals to fly or else there is no effect at all on other members of the
roost. When branches were marked with small pieces of brightly-colored
tape, it was discovered that individual butterflies in a roost almost always
return to the same tree each day for nocturnal roosting. Such specificity
for a given roosting site may imply the use of scent that is laid down and
reinforced on subsequent visits. The problem, however, is very complex
since individuals in a small area can single out their own roosting spots
on different days. Wilson (1968) and Regnier and Law (1968) state
that it is quite possible for different individuals of a species to possess
individual-specific pheromones, resulting from subtle genetic differences,
diet differences or other factors in the environment. Chain-reaction ef-
fects, in which one disturbed individual could initiate flight activity in
other individuals were generally uncommon, although they occasionally
occur. Unlike other known cases of roosting butterflies (e.g., Emmel and
Benson 1971; Clench 1970), due to the relatively large distances among |
members of a roost in Morpho, it is unlikely that mass flight can result
from disturbance directed to a single individual.

Discussion

Adult male butterflies of Morpho amathonte and Morpho granadensis
form sleeping roosts of low densities in tropical rain forests. The low den-
sities of members in these roosts is related to the adult population structure
in these butterflies; adult male populations of Morpho amathonte are very
small and individuals maintain the same individual-specific flight paths
in the habitat. This is probably true for adult Morpho granadensis. Ag-
gressive interactions occur along these flight paths and courtship behavior
has also been observed on them (Young 1971a), and it is suspected that
these flight paths function as territories. Flight paths radiate out from
feeding sites, accumulations of decaying odorous fruit on the forest floor.
Males from roosting aggregations very close to where they feed and both
feeding sites and roosting sites are maintained over long periods of time.
When the feeding site becomes non-functional (with cessation of fruit-fall),
the butterflies disappear and the roosting sites become unoccupied. It
thus appears that in Morpho amathonte, the location and usage of a
roosting site is dependent primarily upon location and usage of a feeding
site. Less is known about the relation of feeding sites and roosting sites
in Morpho granadensis, although roosts are apparently used for long

VoLUME 25, NUMBER 4 229

periods of time. Thus, rather than a roosting site being used throughout
the lifespan of individual adult males, occupancy depends mainly on per-
sistence of feeding sites. Adult Morpho butterflies have been estimated to
live close to 9 months (Young 1971la) and the maximum period of exploi-
tation of certain feeding sites (and thus roosting sites) is 5 months.

The high turnover of adult males in the roost of Morpho granadensis
may be interpreted as the result of predation on adults counterbalanced
by low, but continuous recruitment of newly-emerged adults. In popu-
lations of Morpho amathonte, there is very little recruitment of adults at
feeding sites and there is virtually no mortality of adults from predation
(Young 197la); thus, numbers in roosts tend to remain the same.

Young (197lc) advances a novel hypothesis to account for the dif-
ferential predation by birds on species of butterflies belonging to the
genus Morpho. Under this hypothesis, the apparently high mortality of
adults in Morpho granadensis from predation is explained by the lack of
an effective predator-deterring behavior pattern that is very effective in
adult male populations of Morpho amathonte; adults of the latter species
are successful at escaping from avian predators by attracting them through
marked changes in flight behavior and then frustrating them in relatively
short time. That adult Morpho butterflies are tasteful to avian predators
is shown by (1) the discovery of wing fragments (bearing beak-marks )
of certain species (Morpho granadensis and Morpho peleides) on the
forest floor, under bird perches, (2) observations of attempted attacks by
birds in the field, and (3) records of larval food plants apparently exclude
genera known to contain noxious compounds that make other species
distasteful (Ehrlich and Raven 1965). Larvae of Morpho peleides, a
species closely related to Morpho granadensis (both are in the achilles
group), in El Salvador feed on leaves of Machaerium riparium in the
Leguminoseae (Alberto Muyshondt, V. E. Rudd, personal communica-
tions ); the Leguminoseae are not known for toxic or noxious compounds.
Not only are the present observations a first record of gregarious roosting
in Morpho butterflies, but it is apparently the second recorded case of a
palatable species showing such behavior, as the report of Emmel and
Benson (1971) constitutes the first record for a palatable nymphaline. It
is well known that many species of palatable butterflies are readily taken
as food by caged birds (e.g., Brower and Brower 1964), but less known
is the relationship between palatability and gregarious behavior in trop-
ical butterflies.

Low turnover of adults, in addition to being related to levels of pre-
dation, may be due to a very long developmental time in these butterflies;
for an undetermined species of Morpho in El Salvador, the developmental
time from egg to adult under natural conditions is over 2 months (Senor

230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Alberto Muyshondt, personal communication ). One recent study (Young
1971d) indicated that several species of tropical butterflies belonging to
various genera have very long developmental times. Long developmental
time, low adult population density, and mortality (predation and para-
sitism ) on both adults and immatures may account for the low turnover of
adults in Morpho populations (Young 1971a).

The lack of adult females from the roosts of Morpho butterflies may be
due to undesirable behavioral interactions with males that could occur
there. For Morpho amathonte, it is known that females are relatively rare
at feeding sites where males aggregate, and when they feed there, they
do so when males are absent (Young 197la). Less is known about the
behavior of adult females of Morpho granadensis. Interactions between
sexes do not occur at feeding sites nor at roosting sites in either species;
in Morpho amathonte, courtship takes place along the flight paths of males
(Young 197la). Causal observations at La Selva suggest that adult fe-
males of Morpho granadensis sleep singly on understory vegetation in the
forest; specific locations for sleeping females are highly variable and the
same spot is used for only a few days. Adult females of both species are
probably much more variable in their daily behavior than are males. Fe-
males must continually search for suitable oviposition sites and since they
are probably very long-lived (since males are), they probably remain
fecund throughout the bulk of adult life and this implies a continual search
for oviposition sites. Such a characteristic of their biology, obviously not
shared by males, may account for the lack of similar predictive behavior
in them. Data on the spatial distribution of immatures and how it relates
to the spatial distribution of larval food plants are needed in order to
understand behavior of females in Morpho butterflies.

The absence of females from roosts of males may also be an indication
of a behavioral mechanism that prevents sexual interactions to occur in
places in the habitat where several males may be present at the same time.
The absence of females from nocturnal roosts as well as from feeding sites
of males may prevent severe aggressive interactions that would result in
failure of females to be mated; courtship interactions would be more
successful in aggressive species at places where the chance of multiple-
male encounter would be low. However, such behavior implies that adult
males must have a very effective patrol behavior pattern for females in
such low density populations; this is apparently the case for at least
Morpho amathonte (Young 1971a).

Independent of any gregarious property of roosting in tropical butter-
‘lies, protection to individual members of a roost may result from the
positions of perching. For example, Emmel and Benson (1971) interpret
the perching of roosting Marpesia bernia, a tropical nymphaline, on the

VoLUME 25, NUMBER 4 ell

undersides of leaves as a means of shielding individuals from both rain
and the view of potential avain predators. In the same species, roosting
near the tips of branches may ensure protection against roaming, leaf-
gleaning insectivorous predators such as ants. The roosting of Morpho
butterflies with their wings tightly closed (showing the cryptic under-
sides ) and in vertical positions on the upper surfaces of leaves and near
the tips of branches may also give protection to individual butterflies from
rain and predators. At the time of day when the butterflies arrive at their
roosts, they are very difficult to see when resting (protection against birds),
and their positions on the outer surfaces of hanging leaves gives them a
lot of room to leap out when disturbed (e.g., upon approach of a leaf-
gleaning predator). During heavy rains, I have witnessed various roosting
individuals to go practically untouched by rain resulting largely from their
vertical positions with the wings kept tightly closed.

The repeated occupancy of the same roosting site by the same indi-
viduals over long periods of time suggests that these butterflies have
evolved an efficient means of becoming very familiar with a portion of
their habitat—e.g., it is a form of predictable behavior which such special-
ized behavior is related to protection resulting from coordinated group ac-
tivity is not clear. The lack of close proximity of roosting individuals at a
given roost does not allow for any one individual to “warn” the remaining
members of the aggregation of an approaching danger (predator), and thus
it is very unlikely that gregarious roosting in Morpho butterflies provides
some protection to the group from predation. Rather, it appears that this
behavior is the result of the butterflies aggregating at their feeding sites,
which are very close by (at least for Morpho amathonte). Field observa-
tions support the view that all individuals simultaneously present at a
feeding site will fly away when a human observer walks through the area
(Young 1971la), even though different individuals may be widely sepa-
rated. At feeding sites, gregarious feeding of Morpho butterflies appears
to act as a behavioral mechanism against predatory attacks by resulting
in chain-reaction effects of butterflies flying away. Depending on the
proximity and number of potential nocturnal roosting sites that provide
exposure to morning sunlight, there is a reasonable probability that several
or all adults (since numbers are generally low per feeding site) associated
with a given feeding site will actually roost in the same open spot in the
forest. And as pointed out for certain pierid butterflies (Clench 1970),
the presence of one individual at a roost may attract others to it; such an
attraction, in the case of Morpho, may involve both chemical stimuli and
visual cues. Tethered live individuals or paper models can attract other
living individuals (Young 197la). It thus may be very efficient for all
individuals associated with a given feeding site to select, independently

232 | JOURNAL OF THE LEPIDOPTERISTS SOCIETY

of each other, the closest suitable nocturnal roosting site, so that very little
energy has to be expended in traveling to feeding sites. Such a strategy
appears feasible for at least Morpho amathonte, since adult males of this
species spend a great amount of energy patrolling well-defined flight paths
in search for mates (Young 197la) and since a great deal of time and
energy is put into such reproductive activities, less energy is available
to search for new feeding and new roosting sites on a short-term (e.g.,
day-to-day ) basis. The exploitation of the same feeding sites and roosting
sites on a daily basis allows adult males to partition the bulk of their energy
supply into territorial flying in search of mates.

While similar data are lacking for the daily activity schedule of adult
males of Morpho granadensis, it may be that similar behavior patterns are
present; i.e., the bulk of energy supply is channeled into activities associ-
ated with reproduction and only minimal amounts are used in the selection
of feeding and roosting sites.

Such strategies appear to be of great evolutionary and adaptive signifi-
cance in tropical organisms since population density is generally low in
species that comprise tropical rain forest communities (e.g., Margalef
1968) and under such conditions of low density, a premium is placed
upon highly specialized behavioral patterns that ensure courtship en-
counters. Thus, rather than gregarious roosting in Morpho butterflies
having selective value in terms of predation, a more likely hypothesis,
based on the structure of the roosts and their relation to feeding sites, is
that they are a part of the predictable and efficient overall behavior of
these butterflies that allow for the bulk of the daily energy budget of males
to be used for courtship encounters. The closer spatially that feeding
sites and roosting sites are to one another, the less energy is required to
fly between them. Nocturnal roosting at feeding sites is presumably un-
desirable in that these areas are usually well-shaded from morning sun-
light; roosting in an area exposed to morning sunlight ensures early acti-
vation of the butterflies. For Morpho amathonte, this behavior ensures
early arrival at feeding sites before territorial flying; for Morpho granaden-
sis, although data are lacking, such behavior may ensure initiation of
searches to seek mates, since the bulk of feeding in this species occurs
during late afternoon hours and presumably the food intake on a given
day is put into use the following morning with courtship activities.

Summary

Observations were made on gregarious, nocturnal roosting by adult
males of the large, tropical butterflies, Morpho amathonte centralis and
Morpho granadensis polybaptus at two different sites in Costa Rican trop-
ical rain forests. Individuals of these species roost gregariously in that

VoLUME 25, NUMBER 4 233

several individuals sleep in the same open area in the forest, although they
are close to one another. High fidelity to roost is indicated by occupance
over long periods of time (several months) and turnover of adults is
usually low. Interactions (e.g., aggressiveness) among males does not
occur at roosting sites and females are rare. Although not entirely ruled
out, it is difficult to conceive of such behavior in males of these butterflies
as being a form of collective defense against predators. A more plausible
explanation, based on data on daily activity schedule including feeding
and flying activities for Morpho amathonte, is that such behavior is a by-
product of the gregarious feeding in these butterflies since feeding sites
are located very near roosting sites. As with roosting sites, there is high
fidelity by adult males of Morpho amathonte for feeding sites. Inde-
pendent of the gregarious aspect of this roosting behavior, the type of
position that the butterflies use may give protection against heavy rain
and _ predators.

Acknowledgments

This investigation was conducted while the author was a post-doctoral
associate of a research grant (N.S.F. G-7805) awarded to Dr. Daniel H.
Janzen of The University of Chicago and administered by The Organiza-
tion for Tropical Studies in Costa Rica. The author expresses his thanks
to Dr. Daniel H. Janzen for allowing him to use his facilities in Costa Rica
and to Mr. Jorge R. Campabadal (O.T.S.) for continual logistic support.
Drs. Woodruff W. Benson ( University of Chicago) and Thomas C. Emmel
(University of Florida) assisted in identification of the species studied.

Literature Cited

Brower, L. P. anp J. V. Z. Brower. 1964. Birds, butterflies, and plant poisons: a
study in ecological chemistry. Zoologica 49: 137-159.

Ciencu, H. K. 1970. Communal roosting in Colias and Phoebis (Pieridae). J.
Lep. Soc. 24: 117-120.

CRANE, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hamata
Hewitsen, with special reference to the social use of color. Zoologica 40: 167-197.

CraAng, J. 1957. Imaginal behavior in butterflies of the family Heliconidae: changing
social patterns and irrelevant actions. Zoologica 42: 135-145.

CarpPENTER, G. D. 1933. Acraeine butterflies congregating in a small area for the
night’s rest. Proc. Royal Entomol. Soc. London 6: 71.

Exueuicu, P. R. anp P. H. Raven. 1965. Butterflies and plants: a study in coevolu-
tion. Evolution 18: 586-608.

EMMEL, T. C. AND W. W. Benson. 1971. Survival and gregarious roosting in a
tropical nymphaline butterfly, Marpesia bernia. Evolution (in press).

Levins, R. 1968. Evolution in changing environments. Princeton University Press.

Levins, R. AND R. H. MacArruur. 1967. The maintenance of genetic polymorphism
in a spatially heterogenous environment: variations on a theme by Howard
Levene. Amer. Natur. 100: 585-589.

Marcater, R. 1968. Perspectives in ecological theory. University of Chicago Press.

Minter, R. 1969. Competition and species diversity. In Diversity and Stability in
Ecological Systems, Brookhaven Symposia in Biology, No. 22, pp. 63-70.

934 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Owen, D. F. anp D. O. Cuanter. 1969. Population biology of tropical African
butterflies. Sex ratio and genetic variation in Acraea encedon. J. Zool. Soc.
London 157: 345-874.

Poutton, E. B. 1931. The gregarious sleeping habits of Heliconius charithonia L.
Proc. Royal Entomol. Soc. London 6: 71.

Recnigr, F. E. anp J. H. Law. 1968. Insect pheromones. J. Lipid Research 9:
MULTI.

SLOBODKIN, L .B. AND H. L. Sanpers. 1969. On the contribution of environmental
predictability to species diversity. In Diversity and Stability in Ecological Sys-
tems, Brookhaven Symposia in Biology, No. 22, pp. 82-95.

Witson, E. O. 1968. Chemical systems. In Animal communication, techniques of
study and results of research, T. A. Sebeok (ed.), pp. 75-102, Univ. of Indiana
Press.

Younc, A.M. 197la. Flight and foraging behavior of Morpho butterflies in a trop-
ical rain forest. Ecology (in press).

Younc, A. M. 1971b. Community ecology of some tropical rain forest butterflies.
Amer. Mid]. Natur. (in press).

Younc, A.M. 1971c. An hypothesis on the use of pursuit stimuli by adult Morpho
butterflies as a mechanism of escape from their avian predators. Oecologia (in
press ).

TWO MOSAIC GYNANDROMORPHS OF AUTOMERIS IO
(SATURNIIDAE )

THomas R. MANtLEy!

Bloomsburg State College, Bloomsburg, Pennsylvania

The recognition of a gynandromorph in nature is an exceedingly rare
event. Hessel (1964) described a bilateral gynandromorph of Automeris
io Fabricius taken at Washington, Connecticut. He refers to the capture
of bilateral gynandromorphs of Eacles imperalis Drury taken at Potters-
ville, N. J. (1962) and Callosamia promethea taken at Crown Point, In-
diana (1962) as momentous events in light of the extensively collected
Saturniidae. The appearance of two gynandromorphs in a single season
is unusual, and the data provided by these specimens is extremely valuable
in understanding gene action.

In most Lepidoptera the female is heterogametic. Doncaster (1914),
Xemington (1954), and others have shown the female to have either a
pair of non-homologous sex chromosomes (ZW type) or that one chromo-
some will be lacking (ZO type). The male possesses a homologous pair
(ZZ type).

In A. io. the northern populations are polymorphic; the basic color of

' Research Affiliate in Entomology, Peabody Museum, Yale University, New Haven, Connecticut.

VOLUME 25, NUMBER 4 235

the male is yellow, while the female is a soft, rosy brown. Thus, cells of
the male with ZZ chromosomes are expressed as yellow, while those of the
female with a ZO chromosome complement are rosy brown.

The gynandromorphs described by Hessel (1964) were bilateral, one
side of the body male, the other female. This bilateral condition may have
been caused by a loss of a Z chromosome in the first cell division of the
fertilized egg, or by the development of a binucleate ovum. Such gynan-
dromorphs develop disproportionately in size because the female cells are
larger. Consequently the female side is slightly larger than its male coun-
terpart. Secondary sex characteristics such as form of antenna and the
abdominal claspers, are quite obvious.

The gynandromorphs described in this paper are not bilateral, but
mosaic. They are the first to appear in over 10,000 pupae observed by
this author. Probably the loss or addition of the Z chromosome occurred
at a later state in cleavage causing a disproportionate distribution of cells
with a ZO chromosome expression in gynandromorph no. 19-69. Gynan-
dromorph no. 18-69 suggests another possibility that a ZO cell in its divi-
sion failed to form a cell membrane separating into two ZO cells, leaving
a ZZ cell which produced maie characteristics.

Gynandromorph 19-69 is predominantly male (Figure 1). The head
and antenna are male, the prothorax and mesothorax are composed of male
and female cells, the metathorax is female. The abdomen is male with a
light dispersion of female cells. The upper surface of the right forewing
which is slightly longer than the left, is predominately female with patches
of male cells. The right hind wing is male with a band of female cells
along the anal margin. The eyespot on the right wing is 11 mm by 7 mm
while the left wing eyespot is 9 mm by 7 mm. The difference in size of
the eyespots is caused by the band of female cells on the right wing
whereas the left eyespot is composed entirely of male cells. The forewing
undersides are unique in that identical patterns exist on both wings. A
thin band of female cells is found along the costal margin, with a wide
band of male cells extending from the subcosta radial, area to the medial:
vein. The remainder of the wing is female except a small patch of male
cells on the outer margin of the anal 2 area.

The right hind wing underside is male except for a narrow band of
female cells from the center of the anal 2 area to the inner margin. The
left hind wing underside is female except for male cells extending from the
base along the distance from the base toward the apex.

In specimen 19-69 male cells are in greater proportion than female cells
suggesting that initially the zygote was ZZ but loss of a Z chromosome in
one cell occurred early in the cell cleavage phase causing the female ZO
expression in all future cells developing from it.

236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Fig. 1. Automeris io Fabr. Left, normal male; center, gynandromorph 19-69; right
normal female.

Gynandromorph 18-69 (Figure 2) represents one of the rarest expres-
sions of the characters of A. io, the broken-eye blotch. This feature was
discovered in 1965 among a group of pupae emerging that summer. It
has since been bred to a population where its occurrence can be predicted.
The genetics of the broken-eye blotch will be published later this year.
Its size and coloration suggest it is more female than male. Comparing
its size with normal specimens representing each sex shows this gynan-
dromorph to be considerably larger than most males. This leads to an
interesting speculation. Possibly the fertilized ova was female ZO which
in early cleavage had a cell divide but fail to form a cell membrane di-
viding it into two equal cells. This may have resulted in a cell with a ZZ
chromosome complement which produced only male cells.

The head of this moth is female, as evidenced by the rosy brown color
and antenna. A few isolated patches of male cells are present, notably in
the frons area. The thorax is predominantly female with an occasional
cluster of male cells, the principal one found on the metathorax at the
point of attachment of the right hind wing.

The terga of the abdomen is male with a few small patches of female

VOLUME 25, NuMBER 4 deoafl

Fig. 2. Automeris io Fabr. Left, normal male; center, gynandromorph 18—69 with
broken-eye blotch; right, normal female.

cells. The sterna is a mixture of male and female cells giving a mottled
effect. The terminal, abdominal segment is male on the right side with
a caudal tuft of hair covering the clasper. The left side is female. The
legs are covered with a mixture of male and female colored hair-like
scales.

The color patterns of the forewing uppersides are almost direct op-
posites. The areas on the left side which are male are female on the right
side, and those that are female on the left side are male on the right. The
identation along the margin of the right forewing is due to the smaller
size of the male cells in that area.

The right hind wing on the upperside is male; the left hind wing is
female. The unique nature of this gynandromorph is enhanced by the
multiple expression of the broken-eye blotch. Several complimentary
genes control the expression of the blotch, and these act quantitatively.
The male right forewing reveals the blotch as a single factor expression,
whereas the female left wing shows the blotch in its full expression. The
right wing measures 27 mm, the left wing measures 30 mm.

The forewing underside is female. The right hind wing underside is

238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

female from the costal margin to the medial, vein; the remainder of the
wing is male. The left hind wing underside is predominantly female with
a few narrow lines of male cells, the principal one being a band in the anal
area along the inner margin.

As additional information on the genetics of A. io is unraveled we may
be able to interpret more fully the behavior of cells from zygote to the
mature insect. Controlled breeding is revealing many more unique pat-
terns of this beautiful moth.

Acknowledgments

The author is indebted to Mr. Larry J. Kopp, Klingerstown, Pa., who
reared the larvae; to Dr. James Cole, Bloomsburg State College, who
edited the manuscript, and to Dr. Bradford Sterling, Bloomsburg State
College, for the photography.

Literature Cited

Doncaster, L. 1914. On the relation between chromosomes, sex linked transmission
and sex-determination in Abraxas grossulariata. J. Genet. 4: 1-22.

Hesset, S. A.‘ 1964. A bilateral gynandromorph of Automeris io (Satumiidae) taken
at mercury vapor light in Connecticut. J. Lepid. Soc. 18: 27-31.

RemincTon, C. L. 1954. The genetics of Colias (Lepidoptera). Ady. Genet. 6:
404-407.

NEUTRON IRRADIATION IN ANTHERAEA EUCALYPTI SCOTT
(SATURNIIDAE)

Puittie M. Licuty
Daker Research Center, Stanford, California

Considerable attention has been directed at the entomogenetic effects
of irradiation with 14.1 MeV neutrons. Measurements of neutron induced
anomalies within chromosomes furnishes a biological dosimeter. Moulton
and Meyer (1970) have found in in vivo Drosophila melanogaster that the
frequency of structural abnormalities increases with the radiation ex-
posure level. They demonstrated that the production of dicentrics and
rings corresponds closely to a curvilinear dose response curve. Supportive
evidence for this has been provided by McFee et al. (1970) in in vive
swine leukocytes and Gooch et al. (1964) in human somatic cells.

The present study offers the results obtained from irradiation of in
vitro Antheraea eucalypti Scott tissue cells with 14.1 MeV neutrons.

VoLUME 25, NUMBER 4 239

Taste 1. Chromosome anomalies in in vitro A. eucalypti tissue cells irradiated
with 14.1 MeV neutrons. Values based upon examinations of 200 cells per exposure
level.

Chromatid
Dose Anomalies Deletions Rings Dicentrics
rad N. % (a) (a)

0 2 ) 0 0
100 3 14 1 12
200 6 Al 1! 28
300 BE 78 3 of

Entomogenetic Technique

In a radiation free area, cultures of A. eucalypti tissue cells were reared
from ovaries of diapausing pupae. The moth tissues were incubated at
27° C in an insect tissue culture medium derived by Grace (1962).
Immediately prior to irradiation, samples containing 25 ml of tissue-
containing medium were placed in plastic tubes and situated 8 cm from
the target of a Cockroft-Walton accelerator utilizing the D,T reaction.
Doses of 100, 200, and 300 rads were supplied at a rate of 4 rad/minute.

Following this, the cultures were allowed a 48-hour recovery period.
After a one-hour exposure to 8 x 107 M colchicine, the cells were rinsed
for 15 minutes with distilled water. Cells were fixed for 15 minutes in a
solution of 60 per cent acetic acid—0.1 N hydrochloric acid. Staining with
2 per cent acetic orcein preceded examination of squash preparations for
chromosome structural changes.

Abnormality scoring was obtained by first scanning the coded slides
at low magnification to locate suitable metaphase figures, and then ana-
lyzing them at high power. With the exception of translocations, all struc-
tural aberrations were counted. Metaphase scoring involved 200 cells at
each dosage level.

Results

The neutron irradiation resulted in a pronounced breakage effect, the
amount of which varied with the dosage employed. The frequency and
localization of these abnormalities are summarized in Table I. As ap-
parent from the data, an increase in the dosage level augments the pro-
duction rate of anomalies, but the relationship is not linear. Instead, the
findings more nearly fit a dose square model.

The mean value for one-hit aberrations per cell per rad was 5.63 x 10°
while for dicentrics and rings it was 1.44 x 10°.

At the 300 rad level, evidence of mitotic ruffling was present in about
two per cent of the cells scored. The author believes this to be the first
identification of ruffling induced in moth tissue cells by neutrons.

940 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Acknowledgments

The author would like to extend his deepest appreciation to Dr. Diane
Edwards for her consultation and criticism of the manuscript.

Literature Cited

Goocu, P. C., M. A. BENDER, AND M. L. RANDOLPH. 1964. Chromosome aber-
rations induced in human somatic cells by neutrons. In Biological Effects of Neu-
tron and Protron Irradiations, 1: 325-342. International Atomic Energy Agency.

Grace, T. D. C. 1962. Establishment of four strains of cells from insect tissues
grown in vitro. Nature 195: 788.

McFEE, A. F., M. W. BANNER, AND Mary N. SHERRILL. 1970. Chromosome Aber-
rations in swine leukocytes after in vivo or in vitro exposure to 14 MeV neutrons.
Radiation Research 44: 512-522.

MOULTON, JEAN AND Gar Meyer. 1970. Neutron irradiation in Drosophila melano-
gaster. British Review of Genetics 17: 367-372.

THE “ARROWHEAD BLUE,’ GLAUCOPSYCHE PIASUS
BOISDUVAL (LYCAENIDAE:PLEBEJINAE)

F. MARTIN BROWN
Fountain Valley School, Colorado Springs, Colorado

This started as a short note calling attention to the distribution of
Glaucopsyche piasus Bdy. in Canada. It has developed into a study of
this insect throughout its range. The Arrowhead Blue, like practically
all Plebejinae of North America, was originally placed in the genus Ly-
caena, now reserved for the Coppers. Scudder (1876) created the genus
Phaedrotes with Lycaena catalina Reakirt (1866) as the type species.
Interestingly, Scudder considered that catalina was a synonym of sagit-
tigera Felder (1865) yet designated it the type species. I suspect that he
did this because the type of catalina was known to him and that of sagit-
tigera was in Europe. Edwards (1884) did not accept the new genus, nor
did Skinner (1898), but Dyar did (1902). Placement in Phaedrotes was
followed in general from then (1902) until Nabokov (1945) placed the
Arrowhead Blue in Scolitantides Hubner [1819]. Since then most authors
have followed Nabokov.

The Generic Assignment

Mr. Harry Clench called to my attention the fact that Phaedrotes is in
the tribe Glaucopsychini, and he had demonstrated that Phaedrotes is a
subjective synonym of Glaucopsyche! Huebner’s genus Scolitantides with
Papilio battus | Denis & Schiffermuller] 1775 as type is quite different from

VoLtuMeE 25, NuMBER 4 241

Phaedrotes. lt is closer in appearance and structure to Philotes (sensu
lata) Scudder. Neither Clench nor I can find any structural way to sepa-
rate the Arrowhead Blue from Glaucopsyche. This in spite of marked
pattem differences. The male genitalia, generically “geod” among the
Glaucopsychini, are the same in the two “genera.” The venation is the
same, both have similarly medium-hairy eyes and tousled long-hairy
frons. Both bear androconal scales similar in size and shape with about 12
rows of reticulations each. On piasus these rows converge on the base of
the stalk, on lygdamus they do not. Both utilize legumes as larval hosts.
Scolitantides not only has different male genitalia but the larvae feed on
Sedum and there are no androconia on the wings of the males.

Just enough work was done to fix the place of the Arrowhead Blue. It
shows definitely that a careful world-wide study of the tribe Glauco-
psychini is needed. Clench and I agree that the current concept of Philotes
results in a compound genus. The type species is sonorensis, a Sedum
feeder that lacks androconia (like Scolitantides) whereas the other species
placed with it feed on Oxytheca and Eriogonum and have androconia.

Search of several museum collections demonstrates that Glaucopsyche
piadsus is not common. Personal experience with them is that they are
rather local and never abundant. Perhaps because there are no really
good series to study in any one collection the variation among them has
been overlooked. Eight names have been used with these insects and all
but one of these, daunia Edwards, apply to California specimens.

Study of the specimens available in the Canadian National Collection,
the American Museum of Natural History, the Carnegie Museum, the
Field Museum and my own collection shows that Glaucopsyche piasus
can be divided into two groups of subspecies. For convenience I will call
these the piasus-group and the daunia-group.
piasus-group: lacks a large black spot in Cu:-Cuz in the submargin on the under side

of the hind wing; on the under side of the forewing the spot at the end

of the cell usually is small and linear, and near the middle of the cell
there is a pair of small spots.

daunia-grcup: has a large black spot in Cu:-Cuz and often in Ms-Cu: in the sub-
margin of the under side of the hind wing; on the under side of the
forewing the spot at the end of the cell is large and comma-shaped,
the geminate spots in the middle of the cell often are fused.

The Taxa of the piasus-Group
The earliest name applied to the species is piasus Boisduval (1852).
For many years this name was misapplied to a form of Celestrina from
California. Oberthiir’s figure (1911) shows that the surviving male syn-
type represents the form of Arrowhead Blue found in the lowlands of
central California. On the under side of this form there is very low con-

242, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

trast between the dull greyish ground color and the not much darker
grey patterning (see Figure 2). The type is one of the sort upon which
the arrowhead-shaped white marks are fused into a whitish band. Such
individuals appear in all populations examined.

Another sort of Arrowhead Blue is sagittigera Felder (1865). It is a
form with considerable contrast in the pattern on the under side of the
hind wing (see Figure 5) and with a series of obscure russet spots on the
margin of the upper side of the hind wing of the female. The habitat is
given as “Sonora (Lorquin).” As Brown (1967) pointed out, that means,
according to Boisduval, the vicinity of Los Angeles, California. Sagittigera
has been erroneously placed for many years as a synonym of pidsus piasus.
It is the proper name for the taxon commonly called catalina Reakirt (see
Figures 16-18). Gunder’s slightly aberrant (confluent spots on under
forewing ) gorgonioi belongs to this moiety.

An unrecognized west coast fraction occurs in the northern parts of the
range of the species in California and in Oregon. This is a form on the
underside of which there is considerable contrast, similar to that of sagit-
tigera. These butterflies differ from sagittigera by lacking the russet marks
on the upper side of the hind wings of the female. They are almost uni-
versally confused with piasus piasus from which they differ in degree of
contrast on the under side. I have not seen enough of this variety to name
it and recommend its study to a west coast lepidopterist with sufficient
material. A synopsis of the west coast names follows:

Glaucopsyche piasus piasus (Boisduval), 1852

Ann. Soc. Entomologique de France, 2nd series, 10: 229. Figured by Oberthiir,
1911, Lepid. Compareé, 9: 41, pl. 237, no. 1950 (type). Type locality: “California”
probably the vicinity of San Francisco. Type in U.S.N.M. (see figs. 1-4.) (Ly-
caena piasus )

viaca W. H. Edwards, 1871, Trans. Amer. Ent. Sec., 3: 209. Type locality: “Sierra

Nevada, Cal.” Type in A.M.N.H. (Lycaena viaca)

Range: From the San Bernardino Mountains, California, northward to
at least Butte Falls, Jackson Co., Oregon. Except in southern California,
the subspecies piasus piasus appears from coastal areas up to transitional
forests in the mountains.

Glaucopsyche piasus sagittigera (Felder & Felder), 1865

Reise Novara, Lepidoptera II, p. 281, pl. 35, figs. 20, 21. These figures are poor
but recognizable. Type locality: “Sonora” [vicinity of Los Angeles, Calif.] Type
is in British Museum, Tring, Herts, England. (see figs. 5-6) (Lycaena sagittigera)

catalina Reakirt, 1866. Proc. Acad. Nat. Sci., Philadelphia, 16: 244. Figured by
Strecker (1874) Lepid. Rhop.-Het., pl. 10, fig. 1, 2. These are fair. Type locality:
Los Angeles, California. Type in Field Museum, Chicago, Ill. (see figs. 16-18, the
types.) (Lycaena catalina)

VoLUME 25, NUMBER 4 243

lorquini Behr, 1867, Proc. Calif. Acad. Nat. Sci., 3: 280. Type locality: “California.”
Type destroyed. A letter from Behr to Strecker dated 28.viii.1875 states that lorquini
is a synonym of catalina Reakirt. (Lycaena lorquini )

rhaea Boisduval (1869). Ann. Soc. Entomologique de Belgique 12: 51. Figured
by Oberthiir (1911) Lepid. Compareé 9: 43, pl. 239, nos. 2078 and 2079. Type
locality: vicinity of Los Angeles, California. Type in U.S.N.M. (Lycaena rhaea)

gorgonioi Gunder (1925), Ent. News 36: 4, pl. 1, fig. P (colored). Type locality:

San Gorgonio Mountain, San Bernardino Co., Calif. Type in A.M.N.H. (Phaedrotes

piasus catalina ab. gorgonioi)

Range: Apparently restricted to the coastal regions in the vicinity of
Los Angeles and now in danger of extinction.

The Taxa of the daunia-Group

The daunia-group has much wider range than does the piasus-group
but thus far only one name has been proposed for it. It is found from
northern New Mexico to southern Alberta and west into the Great Basin
and northward to southern British Columbia. Within this extensive range
there occurs some variation.

Material from northern New Mexico (Jemez Springs and Fort Wingate)
tend to show a little russet coloring on the margins of the upper hind wing
of the females. These specimens are in all other respects related to daunia
and not sagittigera. The females of typical daunia from Colorado oc-
casionally show a trace of russet on the upper hind wing. The broad dark
vadum is reduced in the anal region to a submarginal series of dark
patches. These patches of dark scales are linear or lunate and there is a
noticeable lighter area marginad of them.

On the females from British Columbia, Idaho and western Washington,
the vadum is broad, inwardly diffuse and totally lacking any rusty scales,
nor is there any evidence of the broken crenate submarginal line, or its
development, that is common to piasus and daunia. This moiety is recog-
nized below with a subspecific name. _

Glaucopsyche piasus daunia W. H. Edwards, 1871

Trans. Amer. Ent. Soc. 3: 272. Type locality: “Colorado” [Turkey Creek, Jefferson
Co., Colo.] Type is in the Carnegie Museum, Pittsburgh, Pa. (see fig. 7-9). (Ly-
caena daunia)

Range: From northern New Mexico through the Rocky Mountains to
Alberta. The Alberta specimens in the Canadian National Collection, Ot-
tawa, Ontario, are these: Calgary, head of Pine Creek, 3l.v.1915, F. H.
Wolley-Dod, 1 ¢; Cypress Hills, nr. Elkwater, 4500 ft., 13.vii.1951, D. F.
Hardwick, 1 4; Lethbridge, 30.vi.1923, J. McDunnough, | ¢ (illustrated);
Waterton Lakes, 24.vi.1922, C. H. Young, 1 ¢, and 8.vi.1923, J. McDun-
nough, | é.

244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

g ey,

4
g
z
Bes
fod
en
te
8
B

Grrnett

VoLUME 25, NUMBER 4 245

Glaucopsyche piasus toxeuma Brown, new subspecies

Male. Upper side violet blue as on sagittigera, not specular blue as on daunia.
Vadum relatively narrow, inwardly diffuse. Fringes checkered white and black with
the black at the ends of the veins.

Underside: Darker and more contrasty than typical piasus, about as on daunia.
Background about the same shade of grey as on daunia, not as brownish as on sagit-
tigera. Forewing submarginal angular interneural marks poorly defined, especially
in apical area (similar to sagittigera), not as sharply defined as on daunia. Post discal
row, black spots circled with white, essentially the same on all subspecies. The two
cell spots on toxeuma are equally strongly developed and on the whole larger than on
piasus and sagittigera and more like on daunia.

Distal notch of each sagittate white mark on hind wing outlined in black, not as
sharply as on daunia but more like on sagittigera. These marks larger on toxeuma
than on either daunia or sagittigera. In submargin between Cui and Cuz a black spot
capped with a diffuse crescent of rusty scales and then black caret at base of white
sagittate mark. A similar, much less noticeable mark just anterior to anal vein. Oc-
casionally such a mark found in M;-Cu:. These marks absent from piasus and sagit-
tigera and usually more highly developed in daunia than on toxeuma.

Female. Similar coloring to male with broader vadum. Vadum on forewing often
reaching end of cell and diffuse inwardly. On hind wing no evidence of the sub-
marginal sharply defined broken crenate line found on daunia nor diffuse rusty
patches seen on sagittigera.

Under side: As on the males with post discal series of spots often incomplete. In
these cases the minute spots between Rs-M:, Mz-Cu: and Cu:-Cuz disappear.

In addition toxeuma tending to be somewhat larger than specimens of the other
subspecies. |

Holotype. No. 11,470, a male from Garnett Valley, Summerland, B. C., 22.v.1933,
A. N. Gartrell in the Canadian National Collection, Ottawa, Ontario. Radius of left
forewing, 16.7 mm. Allotype: a female from the same series as the holotype but col-
lected on 10.v.1933. Radius of left forewing, 17.0 mm. Paratypes: 9 males and 4
females collected 10—28.v.1933 by either A. N. Gartrell or J. McDunnough. All in the
Canadian National Collection.

The name is a transliteration of the Greek word for “that which is shot,’
in other words, an arrow. The holotype and allotype are shown in Figures
13-15.

Range: South central British Columbia southward in eastern Washing-
ton and Idaho (at least to Pocatello).

Although I have restricted the type series to specimens from the type
locality, the Canadian National Collection contains the following material
assignable to G. piasus toxeuma:

<

Figs. 1-18. Glaucopsyche piasus Bdv. 14, G. piasus piasus Bdy., San Francisco,
Calif., coll. F.M.B.; 5-6, G. piasus sagittigera Felder, Los Angeles, Calif., C.N.C.; 7-9,
G. piasus daunia Edw., Boulder, Colo., C.N.C.; 10, G. piasus toxeuma Brown, Pocatello,
Ida., coll. F.M.B.; 11-12, G. piasus daunia Edw., Lethbridge, Alta., C.N.C.; 13-14,
G. piasus toxeuma Brown, Garnett Valley, B. C., holotype, C.N.C.; 15, G. piasus
toxeuma Brown, Gamett Valley, B. C., allotype, C.N.C.; 16-17, G. piasus catalina
Reakirt, lectotype, Field Mus.; 18, G. piasus catalina Reakirt, paratype female, Field
Mus.

IAB JOURNAL OF THE LEPIDOPTERISTS SOCIETY

BRITISH COLUMBIA: Keremeos, Shingle Creek Rd., 9.vi.1933, A. N. Gartrell,
1 6, and 18.vi.1933, C. B. Barrett, 1 ¢; Okanagan Falls, 15-23.v.1953, J. R. McGillis,
3 6 6; Oliver, 14-15.vi.1953, D. F. Hardwick & J. E. H. Martin, 3 ¢ ¢,1 2; Osoyoos,
19.v.1895, 1 6, 23.v.1895, 1 6, 19-24.vi.1938, J. K. Jacob, 14 ¢¢, 7 2 9, and
15-22.vii.1953, J. R. McGillis, 7 $ 6, 2 9 2; Osoyoos, Anarchist Mt., 7.v.1936, A.
N. Gartrell, 1 ¢@; Penticton, 23.v.1953, J. R. McGillis, 1 ¢; Summerland, Garnett
Valley, 10—28.v.1933, J. McDunnough & A. N. Gartrell, 10 ¢ 6, 5 @ 9, type series
of toxeuma; Vasseau, 14.v.1920, W. B. Anderson, 1 ¢; Vernon, 20.v.1906, Bush-Wilson,
1 ¢, 12.v.1908, 1 6, 22.v.1919, W. B. Anderson, 1 9, and 16.v.1953, D. F. Hardwick
(Sein 10. le Wer, 4 8 6, 5 2.

In addition to the above there is a short series in my collection from
Robson, B. C., collected by N. R. Foxlee on 23-26 May in 1936 and 1937.
Except for these it might be assumed that the subspecies is confined to the
Okanagan Valley in British Columbia. The distributional data for such an
assumption probably is in artifact of collecting. Very definitely there is
a lack of material from both east and west of the Okanagan Valley in the
Canadian National Collection. When that is remedied we will have a
better idea of the range of piasus in Canada.

Literature Cited

(Original descriptions are cited in the text)

Brown, F. M. 1967. Lorquin’s localities “Sonora” and “Utah”. Jour. Lepid. Soc.
21: 271-274.

Dyar, H. G. 1902. A list of North American Lepidoptera and key to the literature
of the Order or Insects. Bull. U.S.N.M. 52.

Epwarps, W. H. 1884. Revised catalogue of the diurnal Lepidoptera of America
north of Mexico. Trans. Amer. Ent. Soc. 11: 245-337.

Nasoxov, V. 1945. Notes on neotropical Plebejinae (Lyaenidae, Lepidoptera )
Psyche 52: 1-65 (esp. p. 3).

ScuppER, S. H. 1876. Synonymic list of the butterflies of North America, north of
Mexico. Bull. Buff. Soc. Nat. Sci. 3: 98-129 (esp. p. 115).

SKINNER, HENRy. 1898. A synonymic Catalogue of the North American Rhopalocera.
Amer. Ent. Soc., Philadelphia.

A RECORD OF EUPHYDRYAS ANICIA (NYMPHALIDAE) IN OKLAHOMA

In June 1959, I took a male of Euphydryas anicia Doubleday at Lawton, Comanche
Co., Oklahoma near Fort Sill, a major artillery training base for the Army. It seems
possible that a pupa or a number of pupae were brought to Fort Sill in some plants a
soldier brought with him or in a shipment of goods to the base. This seems the only
plausible explanation of anicia’s presence in Oklahoma, hundreds of miles from it’s
nearest recorded range.

I have collected in Lawton for eight seasons since 1959, mostly in June, and have
never seen another specimen. This is the first record of the species from east of the
hundredth meridian in Oklahoma that I have knowledge of.

The author would like to thank Mr. F. M. Brown, Fountain Valley School, Colorado
Springs, Colorado, for reading the manuscript and offering his suggestions.

MicHsrEL Toxiver, 1612 Indiana NE, Albuquerque, New Mexico.

VOLUME 25, NUMBER 4 247

REARING TECHNIQUES FOR SPECIES OF SPEYERIA
(NYMPHALIDAE)

S. O. MATTOON
2109 Holly Ave., Chico, California
R. D. Davis
1204 East 13th St., Antioch, California
O. D. SPENCER
935 Ferndale Rd., Lincoln, Nebraska

Widespread interest in North American Speyeria has resulted in ac-
cumulation of considerable taxonomic and ecological information, yet
problems in understanding species limits persist.

Rearing studies would be helpful in clarifying some of these problems,
but until now no successful method of breaking larval diapause or of over-
wintering was known. This paper outlines rearing procedures developed
by the authors during five years of combined research. We have used the
method to rear more than 4,000 specimens, including all the Nearctic
species. Although large scale rearing is described, its fundamentals can
be reduced to the scale desired.

A search of the literature provides little information on rearing Speyeria.
William Henry Edwards (1887) wrote of successfully overwintering larvae
in an ice house. Grey, Moeck, and Evans (1963) reported a method of
breaking diapause by periodic mechanical stimulation over an extended
period. Little was learned concerning the origin of an idea which was
developed into the overwintering procedure described in this paper as
the “block method” of larval storage. The idea came to us through D. V.
McCorkle of Monmouth, Oregon, who along with others explored it as a
means of overwintering.

The Speyerian Life Cycle

A generalized life cycle for Speyeria may be useful in understanding the
complexities of rearing. The one presented is based partly on our obser-
vations and partly on descriptions in literature.

Species of Speyeria are univoltine; adult emergence occurs between
May and September, depending on the population. The males appear
approximately a week before the females, which results from a dispro-
portionate larval growth rate. Females often mate immediately, some-
times on or before their maiden flight. Egg laying begins within days or
weeks depending on differing ovarian maturation rates between species.
Food plants appear to be restricted to the genus Viola (Violaceae) with
one doubtful exception offered by Durden (1965). Oviposition sometimes

948 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

occurs on the food plant, but commonly eggs are placed on other sub-
strates near the host. More than 600 eggs may be deposited by a single
female. The time from oviposition to eclosion under natural conditions
ranges from 12 to 24 days depending on the species. We have noted a
relatively uniform rate of development under laboratory conditions. Eggs
held at temperatures between 24° to 32° C darken within 48 hours if
viable, and hatch in approximately 9 days. After emergence, larvae im-
mediately seek shelter in ground litter and hibernate. All Speyeria larvae
overwinter in a state of diapause, remaining inactive for at least 8 months;
mortality appears to be very high during this period.

Following appropriate stimuli in the spring, feeding starts, and the
larvae grow to maturity in 6 to 10 weeks. The time from pupation to
emergence varies from 7 to 22 days in the laboratory depending on the
species and the temperature to which the pupae are exposed. The average
length of the pupal period in nature is reported to be approximately 14
days, (Weed, 1927; Macy and Shepherd, 1941).

Field Storage and Handling of Live Females

Proper handling permits storage of live females for extended periods
and successful shipping within North America in a condition to produce
an ample supply of eggs. Field collected females should be placed with
their wings closed in glassine envelopes without being stunned or an-
aesthetized. A portable ice chest should be taken where hot, arid con-
ditions will be encountered, or on trips of more than one day. Enveloped
females should be kept in tightly sealed glass jars stored in ice. Lengthy
storage in ice will keep the insects immobile, and is not harmful as long
as desiccation does not occur. The loss of body moisture is a great ship-
ping and storage hazard and may be prevented by the addition of mois-
tened paper towel or the equivalent to the jar. However, adding or ac-
cumulating too much moisture can drown the insects. Also, water can
be accidentally drawn inside from the melting ice especially when a
vacuum is created inside jars as they are cooled or by transporting them
from a high to a lower altitude.

If storage will be longer than 3 days, females should be fed. One initial
feeding will allow them to be carried for several weeks as long as jars are
occasionally opened to replenish the oxygen supply and the cold is main-
tained. Similar success with storage has been obtained in the laboratory
by holding the jars under refrigeration.

Shipment of Living Females

A double box method has proven to be safe and effective for shipping.
The insects (in glassine envelopes) are packed in small loose groups in

VoLUME 25, NUMBER 4 249

dampened absorbent packing material in a durable, crushproof container.
The container is sealed with waterproof packaging tape to help retain
moisture, and is then surrounded with packing material in a slightly larger
container. After packaging, shipments are held under refrigeration until
air mailed. At destination, the females should be fed immediately or the
package again refrigerated.

Adult Feeding Procedure

Females bagged for egg laying should be fed twice daily. The food
solution and feeding procedure is similar to that given by McFarland
(1964). The solution consists of 3 heaping teaspoons of granulated sugar
to 8 ounces of distilled water.

The butterfly, with its wings closed, is placed on absorbent cotton satu-
rated with the solution. Organs of taste in the tarsi of the posterior pair
of legs normally trigger a feeding response (Ford, 1945; Macy and Shepard,
1941; Oldroyd, 1959); however, if the response fails a pin can be inserted
through the proboscis coil extending it into contact with the solution.
Feeding lasts approximately 2 to 3 minutes with old, worn, or egg laying
females. Unproductive individuals may not feed as long.

Adjustable holding devices like the one shown in Figure 1 are con-
venient when numerous females must be fed. In the one shown, a clothes-
pin provided with cardboard grips can be adjusted to the height and angle
necessary for holding any size specimen for feeding. Traces of food
solution should be rinsed away from the insects tarsi and proboscis after
each feeding, otherwise chrystallizing sugar may damage these organs.

Oviposition in Captivity

Following the method used by Grey, Moeck, and Evans (1963), egg
laying can be promoted by enclosing the female in a brown paper bag
exposed to sunlight. In the laboratory an incandescent light source is
used. Size number 8 paper bags containing violet leaves are placed 12”
to 18” from a 100 watt light. The best egg production is obtained when
bags are exposed to a humid environment. Where dryness cannot be
avoided, the paper bags can be enclosed in plastic bags and about 1
tablespoon of water added between them as needed to avoid desiccation
of the females.

Generally most field collected females are gravid, but fresh specimens
of some species must be bagged for many days before oviposition begins.
Females are usually so worn by the end of the egg laying period that
photographic records are necessary to compare progeny and parent.

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VoLUME 25, NUMBER 4 95]

Method of Larval Storage

Newly hatched larvae are immediately removed from the bag and are
housed in small wooden blocks. These blocks, which are approximately
1%” cubes, are preferably made of basswood or other seft lumber. A
hole 2” diameter is drilled with the grain of the wood through the center
of each block to form the storage chamber.

Up to 100 larvae may be housed in an individual storage block. The
storage compartments are closed either by stapling nylon chiffon over the
holes (chiffon is used because it allows ventilation while preventing larval
escape) or by inserting it inside each end by use of plastic compression
rings (Fig. 2). Parakeet banding rings adapt well for closing blocks
because of their ability to expand and contract. Chiffon attachment by
stapling when the blocks are dry will remain tight throughout periods of
expansion and shrinkage resulting from periodic block soaking. Soakings
are mandatory throughout larval storage, as desiccation is fatal to the
larvae. The interval between soakings depends on dryness of storage
conditions, but usually is not longer than a week. Blocks are soaked until
partially wet by absorption in distilled water not deep enough to enter
the storage chamber.

Blocks must be stored at all times under refrigeration at a temperature
just above 0° C, as freezing of the blocks will cause high larval mortality.
The blocks should be arranged in a tray or the refrigerator’s vegetable pan
so that adequate ventilation can pass through the storage chambers. Mold
growth inside damp blocks is a serious problem as mold growing around
larvae will usually cause death. Autoclaving of the blocks before use, and
periodic inspections during storage is necessary. When mold is detected
reblocking must be undertaken.

The overwintering larval condition referred to in this paper and in
literature as “diapause,” may ultimately prove to be quiescence, Through-
out diapause, Speyeria larvae demonstrate an ability to repeatedly awaken
from or return to dormancy in response to the application or removal of
stimuli such as light, heat, and mechanical agitation. The ability to seek
shelter from adverse environmental conditions may have an important
influence in larval survival in nature.

Apparently, a correlation exists between larval metabolism and the agent
controlling diapause. Apparently diapause cannot be permanently ter-
minated until a given amount of stored energy has been expended. The
rate of expenditure seems to regulate termination. Furthermore, the rate
appears to fluctuate with temperature; thus, diapause ends more rapidly
for larvae stored at room temperature (22°-26° C) than for those stored
in the cold (1°-5° C). Carefully controlling the metabolic rate by refrig-
eration to approximate natural habitat temperatures is therefore necessary

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

bo
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777

Ie

Figs. 3, 4. 3, Humidity chamber apparatus for breaking larval diapause (feeding
is promoted by the application of controlled heat, light, and humidity); 4, a bag type
leaf sleeve of nylon chiffon is placed on violet leaf.

to extend diapause to its normal duration. Conversely, the rapid expend-
iture of energy through intense and continued artificial stimulation can
rapidly terminate diapause in all Speyerian species.

Breaking Speyerian Diapause

Figure 3 shows an arrangement developed for breaking Speyerian
diapause. This method employs light, heat, and humidity for stimulus and
protection, and is designed for use at indoor temperatures (22°-26° C).
The equipment consists of a white, enamel plated tray, or other reflective
surface on which is placed a small, clear plastic box or glass petri dish.
Paper towel or filter paper fitted in the bottom of the container is saturated
but not flooded with distilled water. A young tender violet leaf with its
stem held by foam rubber in a 8 X 25 mm water filled specimen vial is
placed in the container. Using a camel hair brush, up to 30 larvae can
be placed in the container. The larvae will partially awaken and contract
into a C-shaped position in response to the handling. An adjustable goose-
neck, hooded lamp using a 60 watt incandescent bulb is positioned ap-
proximately 8 inches from the container shining down on the larvae. After
larvae uncoil and begin to crawl in response to the heat and light stimuli,
they should be transferred to the leaf. A top is placed on the container
to form a miniature escape proof humidity chamber in which the larvae

VOLUME 25, NUMBER 4 ASS)

will not be harmed as long as the towel moisture is maintained. Reflective
insulation material (such as expanded polystyrene) is cut to enclose the
container exposing only the lid. This will substantially reduce condersa-
tion inside the chamber, in which wandering larvae can be trapped and
drowned.

The time required for the first feeding response to occur is highly vari-
able, and as previously stated, appears to depend on how much stored
energy remains to be expended. During this interval the larvae may crawl
extensively with intermittent periods of attempted sleep, often under the
leaf or along the sides of the humidity chamber. The first indication of
feeding is the appearance of small nicks along the margin of the leaf.
Feeding occasionally occurs within 30 minutes, but normally takes from
1 to 3 days or longer. Feeding larvae usually display a gregarious tendency
through the first 2 to 3 instars.

Host Plants

Many North American violets of the genus Viola serve as satisfactory
hosts. Although there appear to be host preferences throughout the genus,
indiscriminate feeding on any available Viola apparently occurs (Macy
and Shepard, 1941, for S. cybele). The only unsuitable violets encountered
are evergreen ornamentals of European origin. Although larvae of most
species will accept an unsuitable host, symptoms possibly indicative of a
nutritional deficiency or a toxic reaction, will develop in varying degrees
depending on the species involved. Some species are quickly killed, while
in others dwarfed adults have been obtained after an abnormally long
development period.

Viola papilionacea Pursh, and the partial albino form V. priceana Pol-
lard have proven to be very successful hosts for all North American Spey-
eria. These large leafed deciduous plants are excellent seed and foliage
producers which withstand repeated defoliation, and can be grown con-
tinuously in greenhouses with only a short winter dormancy.

Rearing Procedures

Larval losses are minimized by allowing completion of the approximate
9-day first instar period in the humidity chamber. They are then trans-
ferred to host plants where second and third instar development continues
on young tender violet leaves in the confinement of a leaf sleeve. Figure
4 shows a nylon chiffon leaf sleeve securely closed by a drawstring which
crimps cotton around the leaf petiole. Violets for sleeving are container
grown for handling convenience. Leaf-sleeving becomes impractical by
late third instar due to increased larval size and food consumption.

As shown in figure 5 the remaining larval development takes place in

254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

a
ye a fe
d | ;
y /
Lo wy

Figs. 5, 6. 5, Nylon chiffon can-sleeve installed on container-grown Viola; 6,
cabinet providing partial temperature and humidity control for storing pupation and
emergence jars in dark.

a nylon chiffon can-sleeve. The wire legs of the sleeve support ring when
pushed into the soil hold the sleeve in position. An elastic band in the
bottom of the sleeve grips the can snugly while access is gained through
the top by means of a draw string.

Violet plants can become infested with several common pest or disease
organisms in addition to harboring hard to detect spiders which can kill
early instar larvae. When pest control becomes necessary, washing the
foliage or hand removal is recommended. Speyeria larvae exhibit a pro-
nounced sensitivity to many commonly used home and garden pesticides,
and any treatment should also include precautionary foliage feeding tests
before plant reuse. Residual or systemic insecticides and those releasing
fumes including household pest strips should be strickly avoided. Soil
in the container may still be contaminated after the plant has proven safe.
A layer of sawdust will prevent soil contact and will collect and help desic-
cate larval droppings for easy disposal.

The feasibility of conducting large scale rearing studies was greatly en-
hanced by learning that the normal more than 3 month larval-pupal devel-
opment period could be significantly reduced. As with most insect devel-
opment, heat is a major factor in shortening the maturation period. In
nature, feeding is slowed or interrupted by the intensity of direct sun-

nine or nighttime cold. Where sleeved cans of larvae are placed under

VoLUME 25, NUMBER 4 2D

continuous incandescent light at a temperature from 26° to 32° C with
moderate humidity, the subdued light within the sleeve seems optimum
for a rapid development of the normally nocturnal larvae.

An extreme example of accelerated development was achieved where
normal males of Speyeria callippe marcaria were obtained in 20 days from
first feeding to adult emergence, but the higher temperature levels needed
are highly favorable for disease development.

Pupal Storage and Adult Emergence

Chrysalids should be stored in separate emergence jars during their
development period. If full-grown larvae are transferred to jars for final
development, the usual indications of oncoming pupation are discontinued
feeding, rapid and continuous wandering, and finally a reddish discolora-
tion to the larval droppings. If pupation occurs in the can-sleeve, masking
tape can be used to easily detach and resuspend chrysalids in the jars
without removing them from their silken attachment.

Newly emerged adults normally undergo nervous body movements
which are apparently a functional part of the wing expansion process.
Three-fourths of the inside circumference of the approximately 2 quart
capacity wide mouth jars, are lined with moisture resistant Dacron curtain
material held in place with “freezer type” masking tape. The lining pro-
vides an adequate foothold surface which reduces wing damage that re-
sults when adults accidently fall during emergence and cannot retain a
hanging position. A gap is left in the lining large enough to lay a res-
taurant dispenser type paper napkin when the jar is on its side. The
napkin, which is kept slightly damp to provide humidity for normal pupal
development, also helps collect larval droppings before pupation and
absorbs the reddish waste material released by the emerging adult.

Addition of water and finally the killing agent can be easily applied to
the napkin with a plastic squeeze bottle through a hole drilled in the jar
lid. Figure 6 shows a jar cabinet designed for large scale rearing. The
storage drawers have removable fronts for easy access to the jars, while
the cabinet provides temperature control and a darkened interior for
emergence.

Acknowledgments

The authors wish to thank research horticulturist Robert L. Smith of
the U. S. Plant Introduction Station, Chico for his continued help and en-
couragement during preparation of the manuscript as well as the use of
his private darkroom facilities. We are especially indebted to Dr. J. A.
Powell, University of California, Berkeley, for reading and suggesting
numerous improvements to the manuscript. T. W. Davies, San Leandro,

956 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

California; D. V. McCorkle, Monmouth, Oregon; Kenneth Tidwell, Salt
Lake City, Utah; J. S. Nordin, Webster, South Dakota and others are
thanked for supplying much of the live material used in our research.

Literature Cited

DurDEN, CHRISTOPHER J. 1965. Speyeria callippe and Artemisia a possible food
plant. Journ. Lepid. Soc. 19: 186-187.

Epwarps, W. H. 1887. The Butterflies of North America. Vol. III. American
Entomological Society, Philadelphia.

Forp, E. B. 1945. Butterflies. The New Naturalist series, Collins Press, London.

Grey, L. P., A. H. Morck anp W. H. Evans. 1963. Notes on overlapping subspecies.
II Segregation in the Speyeria atlantis of the Black Hills. Journ. Lepid. Soc. 17:
129-147.

Macy, R. W., ano H. H. SHeparp. 1941. Butterflies. Univ. Minnesota Press, Min-
neapolis.

McFarianp, N. 1964. Notes on collecting, rearing, and preserving larvae of Mac-
rolepidoptera. Journ. Lepid. Soc. 18: 201-210.

Oxproyp, H. 1958. Collecting, preserving, and studying insects. The MacMillan
Comm Nae

A NOTE ON LETHE ANTHEDON BOREALIS
(SATYRIDAE)

Joun H. Masters

Lemon Street North, North Hudson, Wisconsin

In recent years the genus Lethe Hubner has inspired a good deal of
new interest by North American Lepidopterists; Irwin (1970) has treated
Lethe creola (Skinner ); dosPassos (1969), Shapiro and Carde (1970) and
Carde, Shapiro and Clench (1970) have treated the Lethe eurydice (Jo-
hansson )—Lethe appalachia Chermock complex while Heitzman and
dosPassos (in preparation) are treating the Lethe anthedon Clark—Lethe
portlandia (Fabricius ) complex. As a result of these studies, the number
of recognized species of Lethe in North America is increased from three
(as given by dosPassos 1964) to five. Included are two sets of sibling
species, eurydice-appalachia and creola-portlandia-anthedon, which show
very little morphological divergence but exhibit strong physiological dis-
tinctions which warrant their current designations as separate species. In
view of this, the physiological differences between Lethe anthedon bore-
alis Clark, per my observations in Minnesota, Wisconsin, Ontario and
Manitoba, and nominate L. a. anthedon seem quite pertinent. They should
shed some light on the proper relationship of the two subspecific popu-

VOLUME 25, NUMBER 4 Di

lations and indicate the value in retaining borealis as a subjectively valid
name.

Lethe anthedon borealis was described from Hymers, Ontario by Clark
(1936) as Lethe portlandia borealis in the same paper in which Lethe
portlandia anthedon was described. Borealis has received but little in-
terest, usually having been regarded as a “very weak” subspecies of port-
landia (= anthedon sensu stricto) or as a subjective synonym for anthedon.
The morphological differences, cited by Clark to separate the two sub-
species, are indeed slight; they are as follows:

Dark border on hind wings above narrow and tapering anteriorly; on
the hind wings below the dark band, between the light line bordering
the fourth and fifth spots and the submarginal light line, is little, if at
all, broader than the distance between the submarginal light line and
Pe mMMeMOLt Me WING 2 2 i anthedon Clark
Dark border on hind wings above broader and more uniform, not nar-
rowing appreciably anteriorly; on the hind wings below the dark band
between the light line bordering the fourth and fifth and the submar-
ginal light line is broader, usually much broader, than the distance be-
tween the submarginal light line and the edge of the wing; ground color
below browner and usually more uniform —— borealis Clark

These distinctions are minor and difficult to use in keying out speci-
mens and it is almost necessary to have both populations available for
comparison; however, borealis is not often represented in collections. The
character that I have found most useful in separating populations is the
ground color of the hind wings below; in borealis it is a dull and uniform
brown while in anthedon the background seems to be composed of several
shades of brown and is much brighter. This distinction is especially evi-
dent in looking at the butterflies in series, which avoids comparing in-
dividual differences. I have also found that the morphological distinctions
between the two populations occur on a sharp line between the Transition
and Canadian Life Zones. Munroe (1969) stated that a number of butter-
flies exhibit sharply distinct subspecies on the two sides of a boundary
between two major ecological formations. As examples he cited, among
others, Papilio glaucus canadensis Rothschild & Jordan and Limenitis
arthemis arthemis (Drury) for the Canadian Zone versus Papilio glaucus
glaucus Linnaeus and Limenitis arthemis astyanax (Fabricius) for the
Transition Zone.

After carefully examining the specimens of Lethe anthedon in my own
collection and in the University of Minnesota collection, I placed them,
subjectively, into subspecies borealis or anthedon and plotted their distri-
bution (Fig. 1). The correlation between subspecies and life zones is

258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

PA

Fig. 1. Distribution of examined specimens of Lethe anthedon in Minnesota and
neighboring areas. Circles: Letha a. borealis (Clark). Squares: Lethe a. anthedon
(Clark). Dotted Line: approximate boundary between Canadian (Boreal Region )
and Transition (Austral Region) life zones.

very good; as indicated by the dotted line on the map which traces the
approximate boundary between the Canadian and Transition Zones (per
Roberts, 1936). Nearly every specimen examined could be placed reliably
into one subspecies or the other. Some degree of intergradation was noted
in the character of the dark border on the dorsal hind wing, and to a lesser
extent in the other characters, but in no case was more than one character
involved in intergradation.

The differences in habitat and habits are much more conspicuous.
My observations on Lethe anthedon anthedon have been principally in
southern Minnesota, central Wisconsin, Missouri and Arkansas, but are
in complete accord with Shapiro and Carde (1970) for New York and New
Jersey, Klots (1951) and Edwards (1897). Nominate anthedon is a shade
loving butterfly of deciduous forests. It's foodplants are grasses, Brachy-
elytrum erectum Schreb. (Shapiro & Carde, 1970), Uniola latifolia Michx.
(Heitzman, 1970) and possibly others. The butterflies frequent small glades
in the forest and exhibit strong preferences for flight in the late afternoon,
often flying until dusk. They are infrequently taken at lights which may

VoLUME 25, NUMBER 4 259

indicate that they, like Lethe creola, are subject to occasional nocturnal
flight. The males are very territorial in habit and each male will occupy a
favorite perch at some distance from the perches of his nearest neighbor.
The distance between perches is inversely proportional to the population
density in the particular locality and, if the population density is extremely
high, they may be very close together and, in some cases, even on the same
tree. The perches are almost invariably on a tree trunk, two to four feet off
of the ground, which allows a view of a small glade or opening in the
forest. Territoriality very similar to this has been recorded for a number
of other satyrid butterflies, e.g. Oeneis macounii (Edwards) and Oeneis
jutta (Hubner) (Masters & Sorensen, 1969), and apparently serves the
mating requirements of the butterflies.

Based on my observations, primarily in Rusk County, Wisconsin, Aitkin
and Saint Louis Counties, Minnesota and Riding Mountain, Manitoba,
Lethe anthedon borealis has distinctly different habits and habitats. It
does not occupy the fully wooded environment but prefers a very “open”
wooded environment with lush undergrowth, the mosaic habitat of Shapiro
and Carde (1970). In some cases I have found colonies in localities where
bogs or marshes are gradually giving way to forest and the plant associ-
ations are quite mixed. In other cases I have found them in young seral
forests, composed of birch, aspen and hazelnut growing sparsely with
heavy undergrowths. The foodplant of Lethe anthedon borealis is not
known. One of the recorded foodplants for nominate anthedon, Brachy-
elytrum erectum, occurs at least in part of it’s geographic range (Lakela,
1965), however Uniola latifolia does not. The actual foodplant may prove
to be B. erectum or another grass, but I would not be suprised to leam
that it was a sedge instead since several species of sedge are common in
the borealis habitats.

Like the nominate subspecies, L. a. borealis exhibits a marked preference
for late afternoon flight and may be taken until sunset on warm days. At
Riding Mountain, Manitoba, it is always the last butterfly on the wing
each day and can be taken on overcast days when few or no other butter-
flies are flying.

The greatest behavior difference between borealis and nominate anth-
edon is that borealis exhibits none of the territorial characteristics that are
so pronounced with anthedon and is, in fact, quite gregarious in habit.
Large numbers of borealis are frequently seen congregating together about
a single bush or group of bushes, usually at the edge of a road or a forest
opening. They seem to be quite “amiable” together and the aerial en-
counters of males, that are so common with nominate anthedon and terri-
torial species never occur. Unlike nominate anthedon, they very seldom
perch on tree trunks but prefer perches on low shrubbery, usually less than

260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

a foot off the ground. When alarmed, their favorite tactic is to dodge
deeper into the bush they are on, which makes pursuit by a larger predator
very difficult. Dozens of borealis are sometimes encountered clumped
together while feeding at carrion or excrement or at a shaded, damp spot
in a road. Assemblies of nominate anthedon at damp spots or while
feeding are not uncommon, but the tendencies are not as pronounced and
the numbers involved are not as large.

The range of Lethe anthedon borealis, as far as known, includes: south-
ern Manitoba, west to Riding Mountain; northeastern Minnesota, south
to Aitkin and Carleton Counties; northwest Wisconsin, southeast at least
to Rusk County; and the part of Ontario that is immediately north of Min-
nesota and Lake Superior. It probably occurs further east in Wisconsin
and, quite likely, in the northern peninsula of Michigan. No attempt has
been made to determine the eastern limit in Ontario.

One of the major criticisms leveled at the trinominal and it’s usage in
taxonomy is that the subspecies, as currently defined and used, fails to
distinguish between weakly and strongly differentiated geographic sub-
species and treats them all alike. Descriptions and identifications of
populations are essentially based on visible morphological distinctions and
consequently taxonomy has been strongly oriented in this direction and
populations which show strong morphological divergence have attracted
the most attention. However, we have in Lethe anthedon borealis, a very
good example of very marked physiological (behavioral) differences along
with very weakly developed morphological differences. Differences be-
tween species, subspecies, or any taxonomic category, may be physio-
logical, morphological, or both, and they may be phenotypic or genotypic
in each case. Fortunately physiological differences, which are far more
important in the long run, are usually accompanied by at least minor mor-
phological changes which allows the taxonomist to distinguish and name
the populations exhibiting them. On the other hand, so far as we know,
morphological differences are usually accompanied by at least minor phys-
iological differences; if they weren't, there would be little point in pinning
aname on them. Unfortunately, however, there is not always a correlation.
Remington (1968), for instance, stated that he had under study three
species of butterflies in Connecticut and four in Colorado, each of which
he felt was a pair of widely sympatric and fully speciated entities. He had,
however, delayed formal naming of these species because he had not yet
found [morphological] recognition characters useful for determining mu-
seum specimens.

The North American Lethe constitute a good example of the problems
in relying exclusively on morphological characters for species distinctions.
hrlich (1961) cited Lethe as one of only fourteen North American genera

VoLUME 25, NUMBER 4 261

of butterflies that represented no problem to the taxonomist because
speciation is quite distinct. Since that time field work and behavioral
studies have forced us to increase the number of recognized species from
three to five and we have become aware of classification problems in the
populations of borealis and fumosus Leussler, both of which are still
treated as infraspecific, but with reservations.

The physiological distinctions that I have observed between Lethe
anthedon anthedon and Lethe anthedon borealis are of the same magni-
tude as Shapiro and Carde (1970) found between Lethe eurydice and
Lethe appalachia, and which they used as justification for separating them
into two distinct species. Lethe eurydice and Lethe appalachia are sym-
patric over a wide range, while, as far as is known, Lethe a. anthedon and
Lethe a. borealis are allopatric or nearly so. Thus the retention of them as
subspecific entities is justifiable.

Literature Cited

Carve, R. T., A. M. SHapiro Aanp H. K. Ciencu. 1970. Sibling species in the
eurydice group of Lethe (Lepidoptera: Satyridae). Psyche 77: 70-103.

Crark, A. H. 1936. Notes on the butterflies of the genus Enodia and description
of a new fritillary from Peru. Proc. U. S. Nat. Mus. 83: 251-259.

posPassos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc.
Mem. 1.
1969. Lethe eurydice (Johansson) and Lethe fumosus (Leussler): sibling
species (Lepidoptera: Satyridae). Jour. New York Ent. Soc. 77: 117-122.
Epwarps, W. H. 1897. The butterflies of North America. Vol. 3. Houghton Mif-
flin Co., Boston.

Enruicn, P. R. 1961. Has the biological species concept outlived its usefulness.
Syst. Zoology 10: 167-176.

Herrzman, J. R. 1970. The life history of Amblyscirtes linda (Hesperiidae). Jour.
Research Lepid. 8: 99-104 [1969].

Irwin, R. R. 1970. Notes on Lethe creola (Satyridae), with designation of lecto-
type. Jour. Lepid. Soc. 24: 143-151.

Kors, A. B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Boston.

Laxeta, O. 1965. A flora of northeastern Minnesota. Univ. Minnesota Press, Min-
neapolis.

Masters, J. H. anv J. T. SorRENSEN. 1969. Field observations on forest Oeneis
(Satyridae). Jour. Lepid. Soc. 23: 155-161.

Munroz, E. 1969. Insects of Ontario: Geographical distribution and postglacial
origin. Proc. Ent. Soc. Ont. 99: 43-50.

Remincron, C. L. 1968. A new sibling Papilio from the Rocky Mountains, with
genetic and biological notes (Insecta, Lepidoptera). Postilla No. 119.

Roserts, T. S. 1936. The birds of Minnesota. Univ. Minnesota Press, Min-
neapolis.

Suaprro, A. M. AND R. T. Carve. 1970. Habitat selection and competition in sibling
species of Satyrid butterflies. Evolution 24: 48-54.

262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

DESCRIPTIONS OF LARVAE OF TWO EASTERN SPECIES OF
LYGRIS (GEOMETRIDAE)

W. C. McGurFFIN

Forestry Branch, Department of the Environment,
Ottawa, Ontario

The larvae of seven Canadian species of Lygris were described by the
writer (1958) in Larvae of the Nearctic Larentiinae. Since then two more
species, L. explanata (Walker) and L. molliculata (Walker), have been
reared. The descriptions of these appear here. One other, L. serrataria
Barnes and McDunnough, remains to be studied. A key to known larvae
of Lygris is presented.

Lygris explanata Walker

Cidaria explanata Walker (1862, pp. 1422-1423).

Type locality unknown.

Range: Southern Labrador, Newfoundland, and Nova Scotia, west to Lake of the
Woods and south, according to Forbes (1948), to Massachusetts and New York.

Host: Vaccinium.

Life History: Hibernates as egg; larva free-living (four instars), late May to July
18; pupal period 14 to 18 days; adult late July to August.

Description: I. Head brown; body light brown or yellow, with brown middorsal
stripe; yellow line in upper, and brown line in lower, subdorsal region; wide brown
midventral line. H.W., 0.38 mm; B.L., 3.6 mm; B.W., 0.4 mm. II. Head light brown
with suggestion of brown streaks over lobes. Body light brown with dark brown mid-
dorsal stripe; fine brown line in subdorsal region; brown adventral line; dark brown
midventral line. H.W., 0.54—0.58 mm; B.L., 8-13: mm; B.W., 0.6-0.7 mm. III. Head
light yellow with fine rust-coloured spots arranged in herring-bone pattern on sides of
head, along epicranial stem and on sides of clypeus. Body light brown; middorsal
line dark grey, in an irregular grey stripe; rust line in subdorsal region; adventral line
chocolate; midventral line dark brown. H.W., 0.98-1.00 mm; B.L., 14 mm; B.W.,
1.0-1.1 mm. IV. Head light brown with brown specks arranged in herring-bone pat-
tern along epicranial stem and over parietal lobes. Body light brown; markings dark
grey to reddish brown (Fig. 1). Seta L1 on low tubercle, circled in dark grey.
Thoracic plate concolorous; anal plate concolorous, with dark median furrow. Thoracic
legs light brown; prolegs reddish brown. H.W., 1.67 mm; B.L., 27-35 mm; B.W.,
2.0—2.4 mm.

Remarks: Description based on larvae reared from eggs and collected
from Vaccinium in vicinity of Laniel, Quebec.

Lygris molliculata Walker

Cidaria molliculata Walker (1862, p. 1390).
Type Locality: St. Martin’s Falls, Albany River, Ontario.
Range: Ontario, north and west to the type locality; western Quebec, east to Ste.

Foy; and in the United States, according to Forbes (1948): Minnesota, Pennsylvania
and New York.

Host: Physocarpus opulifolius (L.) Maxim.

VOLUME 25, NUMBER 4 263

PI padded am eth Sel Tre
are ey % ere laeP =o. 8 o

oate ¢

8 Seg Lemire, mai H.
“Seater Cater Ler stig Or
Sn A RR I ARTE nace

om.
sre

; 0 Ne
~

-eNee Le aia

Be SINE a et A ate eee mg rite heme ett de WENN ETS

Figs. 1 and 2. Lygris spp., mature larva, setal and colour patterns on third ab-
dominal segment. 1. L. explanata (Walker); 2. L. molliculata (Walker).

Life History: Larva free-living (five instars), in June; pupal period 14 to 19 days;
adult July.

Description: I. Head light brown; body light brown; middorsal, wide brown line;
narrow brown line in subdorsal region; subventral line wide, brown. H.W., 0.34 mm;
B.L., 5.5 mm; B.W., 0.4 mm. II. Head light brown with brown herring-bone pattern
on lobes. Antenna brown. Body light brown; middorsal stripe grey-brown; brown line
in subdorsal; lateral region greenish; fine grey line in each of subventral and adventral
regions; midventral, very narrow grey line. Legs and plates concolorous. H.W., 0.48—
0.50 mm; B.L., 8 mm; B.W., 0.4 mm. III. Head light grey with much blackish over
lobes; white line over each lobe in line with white line in upper subdorsal region on
thorax. Body light brown; lines grey: middorsal fine, broken; addorsal, as arc on
each anterior abdominal segment; subdorsal, grey, with white line above; subventral,
obscure; adventral, conspicuous; midventral, fine. H.W., 0.70-0.73 mm; B.L., 9-14
mm; B.W., 0.7-0.9 mm. IV. Head whitish, with brown herring-bone pattern on lobes;
whitish on front of head. Antenna light brown, with lines as in last instar. Anal plate
with black middorsal line; prothoracic plate concolorous; legs concolorous, anal with
light stripe along it. H.W., 1.04-1.11 mm; B.L., 18-20 mm; B.W., 1.5 mm. V. Head
light brown, with brown herring-bone pattern over lobes; clypeus often with brown
spot in upper corner, sometimes one in each lower corer as well. Antenna light
brown. Body swollen on mesothorax; light yellow-green to brown, with markings
darker (Fig. 2). Both plates brownish with dark median line. Legs brown, anal leg
with light line along it. H.W., 1.67-1.84 mm; B.L., 25-40 mm; B.W., 1.8-3.0 mm.

Remarks: Description based on larvae collected on west side of Bruce
Peninsula, Ontario.

Apparently some species of Lygris have four-instar larvae (L. explanata)
and some have five-instar larvae (L. molliculata). L. testata L., and L.
xylina Hulst may, apparently, have four or five instars.

Key to Mature Larvae

1. Head horned (parietal lobes attenuate); on Vitis, Parthenocissus, and Epilobium 2

Lleadenonnaleslarvac withvother food: plants 2.2 aS
DVL eae yee ae SxS a en eevee L. diversilineata Hiibner
IPA ACM O LOC ee eae ee ee L. gracilineata Guenée

3. Middorsal line partially connected to addorsal line by transverse bar or dark
patchninivaicimity of DB setae = etree StS aad eee

Middorsal line not connected to addorsal line —.-.------------------ ene usee G

264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

4. Dark oblique posterior to seta D2... eee 5
Dark patch posteriom toi seta: D2) eee L. xylina Hulst
5. Dark transverse bar between D2 setae (Fig. 1) —---.-..- L. explanata Walker
No dark transverse bar between D2 setae _________ L. flavibrunneata McDunnough
6. Setae DI and D2: 0n same dark line 0... 2 ee Z
Setae D1 and D2. not on same dark line 8
f. Middorsalimencomitiraino sees eee L. testata Linnaeus
Middorsal line not continuous (Fig. 2) L. molliculata Walker
8. Oblique dash anterior to seta D1; larvae on Salix and Populus __...---____ 9
No oblique dash anterior to seta D1; larvae on Ribes _____ L. propulsata Walker
9. A continuous dark line through seta SV3 _____.....__._____ L. destinata Moeschler
Little or no dark line through seta SV3 ____.______ L. flavibrunneata McDunnough

Literature Cited

Forses, W. T. M. 1948. Lepidoptera of New York and neighboring states. Pt. 2.
Cornell University Agric. Exp. Sta. Mem. 274.

McGurrin, W. C. 1958. Larvae of the Nearctic Larentiinae (Lepidoptera: Geo-
metridae). Can. Ent. Suppl. 8.

Wacker, F. 1862. List of the specimens of lepidopterous insects in the collection
of the British Museum. Part 25—Geometrites. London.

ON THE GRAMMAR OF THE NAME HELIOTHIS
OCHSENHEIMER (NOCTUIDAE)

GrorRGE C. STEYSKAL

Systematic Entomology Laboratory, Agricultural Research Service,
U. S. Department of Agriculture?

In recent works by Hardwick (1958, 1965, 1970), including extensive
citation of literature, the subfamily name Heliothidinae is used. Investi-
gation into the classical origin of the genus name Heliothis, upon which
that subfamily name is based, reveals that it is an aorist passive participle
of the Greek verb hélio6, meaning to lie in the sun, to bask. This derivation
is given by Treitschke (1826, p. 215) and is confirmed by Ochsenheimer’s
citation in parentheses after Heliothis of the plural nominative form “Heli-
othentes Hiibn.”

As a participle, one of the 2 kinds of verbal adjectives in Greek, there
will be 3 gender forms. These forms and their form in the genitive case,
‘rom which family-group names are formed in the singular number and
trom which names of parasites and other associated organisms may be
tormed in the plural as well as the singular number, are as follows:

' Mailing address: c/o United States National Museum, Washington, D.C. 20560.

)

VOLUME 25, NUMBER 4 265
Masculine Feminine Neuter

Singular Nominative helidtheis heélidtheisa hélidthen
Genitive hélidthentos hélidtheisés hélidthentos

Plural Nominative héliothentes hélidtheisai helidthenta
Genitive hélidthenton helidtheison hélidthenton

In classical transcription into Latin, these forms are:

Singular Nominative heliothis heliothisa heliothen
Genitive heliothentis heliothises heliothentis

Plural Nominative heliothentes heliothisae heliothenta
Genitive heliothenton heliothison heliothenton

Strict application of Article 30.a.i of the International Rules of Zoological
Nomenclature requires treatment of Heliothis as masculine (column 1,
above) and strict application of Article 29 requires use of the stem Heli-
othent- in forming family-group names. This would result in the subfamily
name Heliothentinae. This procedure, according to Hardwick’s citations,
has never been followed.

However, if Article 11.b of the Rules, which states that zoological names
“must be either Latin or Latinized,” be interpreted strictly, we may con-
sider that the complex Greek participial system was not a part of Latin,
except in the case of a few words used as nouns and to be found in Latin
dictionaries. We may then consider Heliothis as declinable in the way
the great majority of Latin nouns in -is are declined, viz.:

Singular Nominative Heliothis
Genitive Heliothis

Plural Nominative Heliothes
Genitive Heliothium

If this be done, the stem used in forming family-group names will be
Helioth- and the subfamily name consequently Heliothinae. At any rate,
there can be no basis for the insertion of -id-.

The matter of gender, however, is something else. Article 30.a.i of the
Rules requires a ruling by the Commission to establish the name as fem-
inine, because it is grammatically clearly masculine and only masculine,
even though ever since Ochsenheimer its usage has been generally as
feminine, probably because the species with adjectival names were origi-
nally described in the genus Noctua. Meigen (1832, p. 224) even changed
the generic name into the feminine from Heliothisa.

Literature Cited

Harpwicx, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera, Noctuidae), with notes on the Heliothidinae.
Canad. Entomol. Suppl. 6.

1965. The com earworm complex. Mem. Entomol. Soc. Canada 40.

1970. A generic revision of the North American Heliothidinae (Lepidop-

tera: Noctuidae). Mem. Entomol. Soc. Canada 73.

966 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

MEIGEN, J. W. 1832. Systematische Beschreibung der europdischen Schmetterlinge.
Aachen, vol. 3.

TREITSCHKE, F. 1826. Die Schmetterlinge von Europa (Fortsetzung des Ochsen-
heimer’schen Werks). Leipzig, vol. 5 (pt. 3).

Editor’s Note: Mr. Steyskal’s paper is an illuminating one and we
are happy to have it for the Journal. I cannot see, however, how Article
11(b) would override Article 29(a) which states: “—if the name of a
type-genus—is a Greek or Latin word—the stem is found by deleting the
case-ending of the appropriate genitive singular.” From a puristic stand-
point, the subfamily name should be the Heliothentinae. This, as well as
the matter of the gender of Heliothis as suggested by Mr. Steyskal, should
be submitted to the Commission for a ruling.

VARIATION IN LARVAL COLOUR PATTERNS OF
ITAME RIBEARIA (GEOMETRIDAE)

W. C. McGuFFIn AND K. BOLTE

Forestry Branch, Canada Department of the Environment,
Ottawa, Ontario

Some geometrid larvae have distinctive colour patterns, others do not;
some species of Itame are good examples of the former (McGuffin 1956).
Dugdale (1961) has shown the importance of infraspecific variation in
colour patterns and has suggested that such variation could offset the
value of coloration for the separation of species. An opportunity to ex-
amine infraspecific variation came in 1968 when a colony of Itame ribearia
(Fitch) was located on an ornamental species of Ribes at Bells Comers,
Ontario.

Of the larvae collected the following were preserved in 70% ethyl al-
cohol: 1 in third instar, 33 in fourth instar and 17 in fifth instar. A small
number was reared; these provided head capsules in second, third, and
fourth instars and information on changes in colour pattern as the larvae
matured.

In the first instar, the head capsule is black; the body is light grey with
a black area surrounding the base of each D seta, a subdorsal grey stripe
of irregular width, and setae L, SV, and V each with a patch of grey at
the base.

In the second instar, the head (3 examined) is light brown with five
black patches, one as a bar at base of clypeus and two on each parietal

VoLUME 25, NUMBER 4 267

lobe, one in ocellar area and one on the upper part of the lobe bearing
setae P and L (Fig. 1); the third abdominal segment (A3), which is
typical of the anterior abdominal segments in this species, is light grey,
suffused with yellow and with black or dark grey patch at base of each
seta, sometimes with grey line connecting patches at base of SD1.

In the third instar, the head (3 examined) is much the same as that in
second instar ( Fig. 2.); A3 is much the same as in second instar with a large
spot bearing setae SD1 and a small spot posterior to former (Fig. 3).

In the fourth instar, five different colour patterns of the head were
noted in an examination of 33 larvae and five head capsules. The basic
pattern of five spots is present with variation in size of the spots on the
parietal lobe. In 17 specimens the patch on the upper part of the lobe
included the base of seta A3; in 3 of the 17 the ocellar patch did not in-
clude seta Al (Fig. 4) but in the remainder it did (Fig. 5). In 16 speci-
mens seta A3 was not included in the patch on the upper parietal lobe
(Fig. 7) and in 7 of these the ocellar patch included seta Al (Fig. 6). In
one specimen the ocellar patch included seta A2 as well as seta Al (Fig.
8). On A3 only two distinctly different patterns were noted. The patches
bearing setae SV1 and SV3 were separate (Fig. 9) in 16 larvae and con-
nected in 9 larvae (Fig. 10).

In the head capsule of the fifth instar the upper parietal patch was
similar in all 17 specimens (Fig. 19) but the ocellar patch in one specimen
(Fig. 20) extended out to include seta Al. Eight variations in the colour
pattern of A3 have been illustrated (Figs. 11-18) and others have been
seen. In all, the subventral stripe is of uneven width and broken or almost
broken between setae L3 and SV4 on the one hand and setae SV1 and
SV3 on the other.

Discussion

This study suggests that the colour pattern of A3 will distinguish ma-
ture (fifth-instar) larvae of I. ribearia from those of I. occiduaria (Pack-
ard), I. evagaria (Hulst) and I. andersoni Swett: the subventral stripe is
broken in I. ribearia but apparently continuous in the other species (Mc-
Guffin 1956). The colour pattern of the head aids in this separation but
by itself is too variable to define the species. In Deilinia and Drepanu-
latrix (McGuffin 1969) the colour patterns of the head were of more
value in separating species than were the colour patterns of the anterior
abdominal segments.

The colour pattern of the head develops from a unicolorous black in the
first instar to a five-spotted one in the second and later instars. On the
other hand the pattern on A3 changes gradually from the first to the last
instar.

968 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

igs. 1-10. Colour patterns of head and third abdominal segment (A3) of Itame
ribearia (Fitch). 1, Second-instar head; 2, third-instar head; 3, third-instar A3; 4-8,
tourth-instar head; 9-10, fourth-instar A3.

VoLUME 25, NUMBER 4

Figs. 11-18. Colour patterns of fifth-instar A3 of Itame ribearia (Fitch).

270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TERS
Seon
De

RTE
fod) oP reres
aR

Figs. 19-20. Colour patterns of fifth-instar head of Itame ribearia (Fitch).

Most of the dark patches on the body of Itame ribearia larvae surround
setae. However, there is one patch with no seta; this patch lies between
setae L1 and D2 on the anterior abdominal segments. It is near the po-
sition of seta SDX2 of Galenara lallata (Hulst) (McGuffin 1967) and
other genera of the Melanolophiini. The dark patch on I. ribearia may
have no relation to the extra seta in the Melanolophiini but its presence
provokes speculations.

Literature Cited

Ducpa.e, J. S. 1961. Larval characters of taxonomic significance of New Zealand
FEnnomines (Lepidoptera: Geometridae). Trans. R. Soc. N. Z. Zool. 1: 215-233.

McGurFin, W. C. 1956. Some larvae of the genus Itame Hibner (Lepidoptera:
Geometridae). Can. Ent. 88: 6-16.

McGurrin, W. C. 1967. Immature stages of some Lepidoptera of Durango,
Mexico. Can. Ent. 99: 1215-1229.

McGurrin, W. C. 1969. Larval head capsule characters for specific identification
in Deilinia and Drepanulatrix (Lepidoptera: Geometridae). Can. Ent. 101: 1228—
IPs

VoLUME 25, NUMBER 4 DLy

PUPAL COLOR IN PAPILIO DEMODOCUS (PAPILIONIDAE) IN
RELATION TO THE SEASON OF THE YEAR

D. F. OWEN

Department of Animal Ecology, University of Lund, Sweden

The pupae of some species of Papilionidae may be either green or
brown. Green pupae match a green leafy background extremely well,
indeed their pattern is suggestive of a curled green leaf, while brown
pupae match a dead brown leaf. Brown pupae are rather more variable
(some are light and others dark brown) than green, which suggests that
~ the color is cryptic as brown vegetation is more variable in color than
green.

The existence of two distinct phenotypes in a population is suggestive
of genetic polymorphism under the control of one pair of alleles with
dominance. But this is evidently not so in the pupae of Papilionidae as
the number of green and brown pupae resulting from a given cross does
not fit any known segregation ratio. A typical result is for almost all the
pupae to be one color and one or two the other color. In Papilio demo-
docus green X green crosses have given some brown pupae and brown X
brown crosses some green pupae. These results cannot be explained by
the existence of a pair of alleles with dominance as either green or brown
would have to be homozygous recessives and would give only one pupal
color among the offspring. It therefore appears that pupal color is en-
vironmentally determined, although there may be a genetic component
to the capacity to produce a pupa of a particular color.

One possibility is that green pupae tend to be produced when the larvae
pupate on a green background and brown pupae when on a brown back-
ground. There is some evidence of this which will be discussed later, but
in addition there is evidence of a seasonal cycle in the production of green
and brown pupae.

On the assumption that green and brown pupae are environmentally
determined an experiment was planned to determine if the season of the
year affected the frequency of the two pupal colors. There are a priori
reasons for supposing that green pupae would be advantageous in a
tropical wet season and brown pupae in a dry season. It was possible to
perform the experiment in both West Africa and East Africa at localities
that are quite different in climate. The species chosen for investigation,
Papilio demodocus, is a common African butterfly. The larvae now feed
on Citrus, which is an introduced plant in Africa, but some larvae may
still be found on the presumed original foodplants, various wild species of
Rutaceae.

272, JOURNAL OF THE LEPIDOPTERISTS SOCIETY

TABLE 1. Mean monthly rainfall (in mm) at Kampala, Uganda, and Freetown,
Sierra Leone.

J BY M A M J J A S O N D

Kampala 64 53 139 174 95 61 ol ie 98° Wiser 96
Freetown 6 1 iM 60 180 382 3803 849 600) 250 aig, 25

Methods

First and second instar larvae were collected in the wild on Citrus and
reared in small glass jars, one or two larvae to each jar. When full grown
they were allowed to pupate on the side of the jar, but not on the food-
plant. The jars were placed indoors and the larvae were reared under
normal conditions of temperature, but a very high humidity (near 100% )
was generated and maintained inside the closed jars by the larvae and the
plants. Larvae were collected and reared in all months of the year. The
experiment was conducted in two parts, the first at Kampala, Uganda, in
1964-65, and the second at Freetown, Sierra Leone, in 1968-69. Both
these localities have an equatorial climate, but differ in that in Uganda
there is some rain in all months of the year with two seasonal peaks, while
at Freetown there is a very large single peak of rainfall and a rather severe
dry season. The mean monthly rainfall figures (ten-year averages) are
shown in Table 1. The most important ecological difference between the
two localities which results from the seasonal distribution of rainfall is
that Kampala is relatively green all the year round while at Freetown the
dry season causes a considerable dying back of the vegetation and the
environment is for several months quite brown.

In addition to the main experiment some larvae were reared in the dark
in blackened jars and some were reared at high densities, but the results
are ambiguous and are not discussed further.

Pupal Color and the Season of the Year

In Table 2 the pupae obtained during the experiments are divided into
two groups corresponding to the six wettest and six driest months at Kam-
pala and to the wet and dry season at Freetown. Just over 66 per cent of
the pupae at Kampala were green and there is no seasonal difference. But
at Freetown green pupae were significantly more frequent in the wet
season and brown in the dry season (,? = 32.4, P< 0.001). In the dry
season just over 39 per cent were green while in the wet season nearly
twice as many were green. Since the larvae were forced to pupate on the
sides of the glass jars there is no question of matching the background and

‘ appears that there is a built-in seasonal cycle in the frequency of green

VOLUME 25, NUMBER 4 Da

TaBLE 2. Relative frequency of green and brown pupae in Papilio demodocus reared
under controlled conditions at Kampala, Uganda, and Freetown, Sierra Leone.

green brown % green
Kampala
Six driest months (Dec.-Feb., May-Jul. ) 28 14 66.7
Six wettest months (Mar.-Apr., Aug.-Nov. ) 43 WO) 66.2
Freetown
Dry season (Nov.-Apr. ) 48 74 39.3
Wet season (May-Oct. ) 89 28 Gell

and brown pupae at Freetown, which, however, does not occur at
Kampala.

In November 1970 at Freetown 16 larvae were reared in a cage con-
taining a pot of growing Citrus. Six pupated on the plant and all formed
green pupae, while the remainder pupated on the wooden lid of the cage
and produced brown pupae. The sample is rather small but the result is
suggestive that pupal color is correlated with the background on which
the larvae pupate.

Discussion

The results obtained suggest that the color of Papilio demodocus pupae
is adapted to the background color on which pupation takes place, and
that in an area where there is a conspicuous seasonal change in the back-
ground color there is a corresponding cycle in the frequency of green and
brown pupae. Exactly how the cycle is generated is obscure. The di-
morphism appears to be largely environmentally controlled but this of
course does not exclude the possibility that there are genes that affect
larval behavior. It is possible to postulate a switch mechanism which acts
on the larva in response to the background on which it is pupating, but
in addition there may at Freetown be a seasonal cycle in larval behavior
such that more seek out green backgrounds in the wet season than in the
dry. On the other hand there are more green backgrounds available in the
wet season and thus the probability of a larva pupating on green is
increased.

Further investigation is required, especially as other species of Papil-
ionidae also produce green and brown pupae (at Freetown, Papilio
nireus and Graphium policenes), while there are evidently species in
which the dimorphism is absent; thus all the pupae of Papilio dardanus at
Freetown seem to be green.

274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Summary

The green and brown alternative pupal colors in the butterfly, Papilio
demodocus, appear to be environmentally determined. Larvae reared
under controlled conditions in Uganda produced about two-thirds green
and one third brown throughout the year, but in Sierra Leone green pupae
were more frequent in the wet season and brown in the dry. The dif-
ference between the two localities is correlated with differences in the
seasonal distribution of rainfall and its effect on green and brown back-
ground colors. There is some evidence that green pupae are formed on
green backgrounds and brown pupae on non-green backgrounds.

THE LIFE HISTORY OF SCHINIA LIGEAE (NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute, Canada Department of Agriculture,
Ottawa, Ontario

Schinia ligeae (Smith, 1893, p. 331) feeds in the larval stage on the
blossoms of the Mojave Aster, Machaeranthera tortifolius (Gray) (Figs.
2, 4). According to Munz (1963), tortifolius occurs on the Mojave and
northern Colorado deserts of southern California and is distributed from
there eastward to southwestern Utah and western Arizona.

Specimens of Schinia ligeae in the Canadian National Collection were
taken in various localities on the northern Colorado Desert and the Mojave
Desert except for one specimen taken in the Valley of Fire in southern
Nevada and another at Ehrenberg, Arizona. The distribution of the moth,
therefore, may well correspond with the distribution of its host plant.

Schinia ligeae is a spring-flying species and its flight is co-ordinated with
the early blossom period of its host plant. Specimens examined were taken
between the middle of March and the end of April.

Behaviour

Schinia ligeae is a predominantly nocturnal species and the eggs are
deposited at night in the heads of the Mojave Aster. The eggs are inserted
among the florets from the upper surface of the head, usually at a stage
when the individual florets of the blossom have not as yet opened. On
the basis of the few eggs observed throughout the incubation period,
eclosion usually occurs on the fifth day after deposition. The newly
hatched larva bores into an adjacent floret and feeds within it for the
duration of the first stadium. The second-stadium larva feeds in the head

VoLUME 25, NuMBER 4 215

Figs. 1-6. Schinia ligeae (Smith), its habitat and food plant. 1, Adult, Desert
Hot Springs, Calif.; 2, Fan Hill Wash near Desert Hot Springs where eggs and larvae
were found abundantly in the heads of Mojave Aster; 3, ventral aspect of pupae; 4,
food plant, Machaeranthera tortifolius (Gray); 5, 6, fourth-stadium larvae.

from a position between the florets. Early in the fourth stadium the
larva may leave the first head and bore into a second, which it attacks
from the top. The larva remains concealed within the aster heads through-
out its period of feeding. The fully fed larva makes its way to the ground
and tunnels into the soil to pupate. All individually reared larvae matured
in four stadia.

Description of Stages

The few gravid females taken in southem California did not oviposit
readily in captivity and the developmental data recorded here were based

276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 7, 8. Apical abdominal segments of pupa. 7, Ventral; 8, right lateral.

largely on individuals dissected as eggs from the heads of Mojave Aster
at Fan Hill Wash near Desert Hot Springs, California. Larvae were reared
individually at room temperature using the techniques outlined by Hard-
wick (1958). The estimate of variability following the mean for various
values is the standard deviation.

Adult (Fig. 1). Head and thorax pale fawn-grey. Abdomen fawn-grey or oc-
casionally smoky-grey. Forewing pale grey marked with light fawn. Transverse an-
terior line white, usually smooth, acutely excurved. Basal space light fawn, usually
becoming paler toward base of wing. Transverse posterior line white, inconspicuously
scalloped between veins, shallowly excurved around cell, then straight to inner
margin. Median space whitish-grey. Orbicular and reniform spots absent. Sub-
terminal space narrow, concolorous with basal space. Terminal space cream. Often
a series of indistinct intervenal dashes at margin of wing. Fringe concolorous with
terminal space. Hind wing light to dark smoky-brown. An indistinct discal lunule and
outer-marginal band sometimes evident. Fringe white with a yellowish basal line.
Underside of forewing with basal and median areas uniform chocolate-brown; outer
and. apical areas of wing dull cream. Hind wing dull cream, usually with an evanescent
discal lunule.

Expanse: 26.9 + 1.6 mm (62 specimens ).

Egg. Pale cream when deposited, becoming suffused with pink on day after
deposition. Pink darkening to an orange-brown during next few days; anterior half of
egg darker than posterior half. Egg becoming grey, then head capsule becoming
visible through chorion a few hours before hatching.

Dimensions of egg: length, 1.55 + 0.08 mm; diameter, 0.87 + 0.06 mm (25 eggs).

First-Stadium Larva. Head varying from medium orange-brown through smoky-
brown to blackish-brown. Prothoracic shield smoky-fawn to dark smoky-brown.
Suranal shield similar in colour but usually paler. Trunk translucent yellowish-grey
in newly hatched larva with purplish gut showing through; older larvae cream to
light yellow. Spiracles with dark-brown rims.

Head width: 0.518 + 0.017 mm (25 larvae).

Duration of stadium: 4.5 + 0.6 days (39 larvae).

Second-Stadium Larva. Head light to medium orange-brown, often mottled

VoLuME 25, NuMBER 4 207

dorsally with smoky-brown. Prothoracic shield varying from light orange-brown to
fawn; variably suffused with dark smoky-brown, often heavily so; a pale median line
often evident on shield. Suranal shield paler than prothoracic shield. Trunk varying
from cream to light yellow. Spiracles with medium- or dark-brown rims. Thoracic
legs fawn, variably suffused with smoky-brown.

Head width: 0.864 + 0.031 mm (25 larvae).

Duration of stadium: 3.4 + 0.7 days (39 larvae).

Third-Stadium Larva. Head bright orange-brown, suffused and mottled with
smoky-brown. Prothoracic shield black, margined with fawn or smoky-brown, and
with a narrow median line of pale grey or light cream. Suranal shield varying from
fawn-grey to orange-brown. Trunk greyish-cream, without maculation except for in-
creased transparency over heart. Spiracles with black rims. Thoracic legs varying
from light fawn to pale orange-brown and suffused with smoky-brown.

Head width: 1.41 + 0.07 mm (25 larvae).

Duration of stadium: 3.7 + 0.7 days (39 larvae).

Fourth-Stadium Larva (Figs. 5, 6). Head bright orange, almost indiscernibly
mottled with light orange-brown. Prothoracic shield varying from fawn to light
orange-brown, with a black or smoky-brown submarginal patch on either side and
with a narrow, pale yellow, median line. Suranal shield light orange-brown, weakly
mottled with smoky-brown. Trunk various shades of grey, usually slate-grey on
entering stadium but becoming paler with increasing size; trunk assuming a dull red-
dish tone on dorsum as larva approaches the prepupal phase. Spiracles with black
rims. Thoracic legs varying from dull yellow to light orange-brown.

Head width: 2.09 + 0.12 mm (8 larvae).

Duration of stadium: 6.3 + 1.4 days (39 larvae).

Pupa (Figs. 3, 7, 8). Mahogany-brown, suffused with olive on proboscis and
thoracic appendages. Spiracles on segments 2, 3 and 4 borne in shallow oval depres-
sions; spiracles on segments 5, 6 and 7 borne in noticeably deeper pits. Spiracular
sclerites moderately broad. Anterior marginal area on each of abdominal segments 5,
6 and 7 strongly sclerotized and raised into a rounded, prominently pitted ridge.
Proboscis terminating a short distance anterior to apexes of wings. Cremaster usually
consisting of two rather short setae borne at the narrowly rounded apex of the tenth
abdominal segment; a few pupae with a short slender seta lateral to each of the
median pair.

Length from anterior end to posterior margin of fourth abdominal segment: 8.9 +
0.4 mm (24 pupae).

Acknowledgments

I appreciate the assistance of Mr. John E. H. Martin, Entomology Re-
search Institute, in providing the photographs for this paper, and of my
associate Mr. Eric Rockburne in measuring larval structures and in draw-
ing the cremaster area of the pupa.

Literature Cited

Harpwickx, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

Munz, P. A. 1963. A California flora. University of California Press, Berkeley.

SmirH, J. B. 1893. Descriptions of Noctuidae from the Death Valley. Insect Life
5: 328-334.

278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

INTER-SPECIFIC HYBRIDIZATION BETWEEN LIMENITIS
ARTHEMIS ASTYANAX AND L. ARCHIPPUS (NYMPHALIDAE)

AUSTIN bl by Amen
University of Maryland Baltimore County, Catonsville, Maryland
and.
JosepH C. GREENFIELD, JR.
Duke University Medical Center, Durham, North Carolina

The Nearctic genus Limenitis (Nymphalidae) contains five common,
geographically widespread forms, all of which are polytypic, and exhibit
tendencies toward hybridization (Edwards, 1879; Scudder, 1889; Field,
1904; Gunder, 1934; Remington, 1958, 1968; Gage, 1970). Four of the
forms are mainly allopatric in their distributions, occupying adjacent re-
gions, and coming in contact only along certain margins of their ranges
(Hovanitz, 1949). Included among these are two conspecific eastern
forms: the banded purple (L. arthemis arthemis Drury) and the red-
spotted purple (L. arthemis astyanax Fabricius ), an unbanded mimic of
the blue swallowtail (Battus philenor L.). In addition, there are two
western disruptively banded species: Weidemeyer’s admiral (L. weide-
meyeriti Edwards) and Lorquin’s admiral (L. lorquini Boisduval).

These four forms are closely allied, and conform well to Mayr’s (1963)
definition of a “super-species.” The two subspecific eastern butterflies
exhibit “free-interbreeding” and complete intergradation within the north-
eastern United States and southern Ontario (Edwards, 1877; Field, 1910;
Hovanitz, 1949; Platt and Brower, 1968; Remington, 1968; Platt, Frearson,
and Graves, 1970), whereas, the two western species exhibit “suturing”
and “intense” interbreeding in certain restricted localities, often associated
with mountain passes (Brown, 1934; Perkins and Perkins, 1966; Perkins
and Perkins, 1967; Remington, 1968).

The fifth form is the predominantly orange-colored Viceroy (L. arch-
ippus Cramer). It is broadly sympatric with all four of the others and
represents a distinct species having: a) evolved a mimetic color-pattern
closely resembling the unpalatable monarch (Danaus plexippus L.), b)
possessing highly modified male claspers (Scudder, 1889; Nakahara,
1924; Chermock, 1950; Platt, Frearson, and Graves, 1970), and c) being,
in part, at least, ecologically isolated from the others, preferring open
marshy meadows to woods-meadow ecotones and woodland glades.

The purposes of this report are: first, to document the occurrence of a
recently collected wild hybrid between the two mimetic species, L. a.
astyanax and L. archippus; second, to review previously known records of
such inter-specific wild hybrids in order to verify the scarcity and wide
geographic distribution of such specimens; and, third, to present a pre-

VOLUME 25, NUMBER 4 279

lem

Fig. 1. Wild-caught hybrid male (form rubidus Strecker) from Durham, N. C.
with parental species; top row dorsal, bottom row ventral. Left, Limenitis archippus;
middle, hybrid form rubidus; right, Limenitis arthemis astyanax.

liminary report of recent laboratory crosses which confirm the hybrid
nature of these intermediate “suspected” wild hybrids.

The new hybrid specimen (Figure 1) is a male, collected on Highway
751, eight miles south of Durham, Durham County, North Carolina, on
October 10, 1970 by J. C. Greenfield, Jr.1 It can be referred to hybrid form
rubidus Strecker, in that its basic ground color is orange like that of
archippus but the dorsal surfaces of the forewings are darkly pigmented,
whereas, the hind wings possess large red-orange marginal spots. Ven-
trally, both the proximal and marginal red-orange spotting and remnants
of the double row of marginal iridescent lunules characteristic of astyanax
are present. Both parental species fly commonly in the fields and woods
surrounding the vicinity where the specimen was secured.

Other known records of wild hybrids between butterflies of the L. a.
arthemis-astyanax complex and L. archippus are given in Table 1. By
reviewing the literature, and corresponding with museum curators and
Society members, a total of eight records of wild arthemis x archippus
hybrids (form arthechippus Scudder), and 12 previous reports of wild-
caught rubidus have been found. The wild specimens are widely distrib-
uted, but exceedingly rare, those reported in Table 1 having been collected
from 1872-1970. Insofar as is known, all wild specimens collected to date
have been males.

1 This specimen has been donated to the collection of the American Museum of Natural History
in New York.

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

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bien

Fig. 2. Lab-bred Fi L. arthemis astyanax x L. archippus hybrids; top row dorsal,
bottom row ventral. Left, light (archippus-like) form; right, dark (astyanax-like) form.
Specimens bred from Maryland stocks in January, 1970.

Proof of the hybrid nature of rubidus requires experimental crosses
between the two parental species. As noted in Table 1, there is at least
one record of astyanax and archippus having been collected in copula in
the natural environment. Such hybrid crosses recently have been made by
Platt (Figure 2) by hand-pairing the insects (Platt, 1969), and earlier ones
are known to have been done at Yale University (C. L. Remington and
R. W. Pease Jr. personal communication). To date, seven fertile crosses
have been obtained by Platt. Since these data will be reported in greater
detail later, only a brief report of the findings will be included here.

Five crosses between astyanax females and archippus males have pro-
duced a total of 52 F,; male progeny. Two crosses between archippus
females and astyanax males have yielded 78 F; males, for a combined total
of 130 F, males. As shown in Figure 2, the inter-specific hybrids occur
in both light (more archippus-like ) and dark (more astyanax-like) morphs,

282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

the wild-caught specimen described above being of the darker variety
(Figure 1). In contrast to arthechippus hybrids, all rubidus entirely lack
the partial white band markings on their dorsal surfaces. However, some,
but not all, have traces of the white band persisting in the costal regions
of the forewing, as does the recent wild-caught specimen. Such white
markings are more fully expressed in archippus, but similar markings
also are found in certain individuals of astyanax (Clark and Clark, 1951;
Platt and Brower, 1968).

The complete documentation of rubidus as an interspecific hybrid is
shown by the fact that all bred F, specimens obtained to date are males.
Complete heterogametic (female) inviability is encountered when the
two full species are hybridized, in accordance with Haldane’s Rule (Steb-
bins, 1958; Bowden, 1966).

However, recently the rubidus hybrids have been backcrossed success-
fully to both astyanax and arthemis females, and to archippus females, as
well, yielding four viable broods having a total of 34 male and 12 female
progeny (46 in all). Recovery of some of the females is noteworthy,
although the sex ratios are still biased in favor of males (X?, = 10.52,
P <0.1). Only three males have been obtained so far in two backcrosses
to archippus females, and breeding experiments are continuing.

Nevertheless, these preliminary broods demonstrate that the F; male
hybrids are fertile in backcrosses to the parent females. Genetic and
phenotypic segregation also is apparent in these crosses, all combinations
of which have yielded “parent-like” and “hybrid-like” individuals.

Environmental selection probably is operating against the rare natu-
rally occurring hybrids. Since the two parental species are considered to
be Batesian mimics of two totally different unpalatable models, such
intermediate morphs become exceedingly poor mimics of either one. The
total female inviability encountered in F, crosses means that the wild
males must have to breed with parental females, if at all. Platt, Frearson,
and Graves (1970) have shown that arthechippus males possess valvae
intermediate in shape between those of the two parent species; the same
also undoubtedly is true of rubidus males. Consequently, sexual selection
and mate choice, in which coloration and courtship behavior likewise are
important, would not seem to be favoring the male hybrids.

In conclusion, the rare wild hybrid form rubidus Strecker represents a
true inter-specific F,; hybrid arising from “stray” matings between two
closely related, but distinct mimetic species, L. a. astyanax and L. arch-
ippus. As such, it illustrates well the breakdown of Batesian mimicry in the

iatural environment, and the selective elimination of an unfit phenotype.

VoLUME 25, NUMBER 4 283

Acknowledgments

We are indebted to Dr. A. B. Klots and Dr. F. H. Rindge of the Amer-
ican Museum of Natural History for confirmation of the identity of the
wild hybrid specimen, and for providing certain information given in
Table 1. The assistance of Society members who have corresponded with
Dr. Platt about hybridization in Limenitis also is gratefully acknowledged.

Literature Cited

BowveNn, S. R. 1966. “Sex-ratio” in Pieris hybrids. J. Lepid. Soc. 20: 189-196.

Brown, C. 1934. Notes on Basilarchia lorquini Bdy., form fridayi Gun. (Lepid.:
Nymphalidae). Entomol. News 45: 205-206.

CuHerMock, R. L. 1950. A generic revision of the Limenitini of the world. Amer.
Midl. Nat. 43: 513-569.

Criark, A. H., anp L. H. Crarx. 1951. The butterflies of Virginia. Smiths. Misc.
Coll. 116: 1-239.

Epwarps, W. H. 1877. Notes on Limenitis proserpina and arthemis. Canad. Ent.
9: 114.

Epwarops, W. H. 1879. Butterflies of North America. Vol. 1: 111-147. (Privately
printed ).

Epwarps, W. H. 1882. Description of new species of butterflies found in the
United States. Papilio 2: 45—49.

Fretp, W. L. W. 1904. Problems in the genus Basilarchia. Psyche 11: 1-6.

Fretp, W. L. W. 1910. The offspring of a captured female Basilarchia. Psyche
Ae VS —117.

Frep, W. L. W. 1914. Hybrid butterflies of the genus Basilarchia. Psyche 21:
115-117.

Gacr, E. V. 1970. A record of a naturally occurring Liminetis hybrid. J. Lepid.
Soc. 24: 270-271.

Grey, L. P. 1968. (No title) In North American Annual Summary. News Lepid.
Soe. No. 3, p. 19.

Gunver, J.D. 1934. A check list revision of the Genus Basilarchia Scud. ( Lepid.:
Rhopalocera). Canad. Ent. 66: 39-48.

Hovanitz, W. 1949. Increased variability in populations following natural hy-
bridization. In Jepsen, G. L., E. Mayr, and G. G. Simpson (Eds.). Genetics,
paleontology, and evolution. Princeton Univ. Press, pp. 339-355.

Kuiots, A.B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Boston,
pp. 115-116.

Mayr, E. 1963. Animal species and evolution. Belknap Press of Harvard Univ.
Press, Cambridge, Mass.

Monroe, B. L. 1953. A hybrid Limenitis. Lepid. News 7: 53.

Naxanara, W. 1924. A revision of the genus Basilarchia (Rhopalocera: Nymphal-
idae). Bull. Brooklyn Ent. Soc. 19: 166-180.

Newcoms, H. H. 1907. Description of a new variety of Limenitis ursula. Psyche
14: 89-91.

Perkins, E. M. ann S. F. Perkins. 1966. A review of the Limenitis lorquini com-
plex (Nymphalidae). J. Lepid. Soc. 20: 172-176.

Perkins, S. F. anp E. M. Perkins, Jr. 1967. Revision of the Limenitis weide-
meyerii complex, with description of a new subspecies (Nymphalidae). J.
Lepid. Soc. 21: 213-234.

Puatr, A. P. 1969. A simple technique for hand-pairing Limenitis butterflies
(Nymphalidae). J. Lepid. Soc. 23: 109-112.

Piatt, A. P. anp L. P. Brower. 1968. Mimetic versus disruptive coloration in

284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

intergrading populations of Limenitis arthemis and astyanax butterflies. Evolution
22: 699-718.

Puatt, A. P., S. D. FREARSON, AND P. N. Graves. 1970. Statistical comparisons of
valval structure within and between populations of North American Limenitis
(Nymphalidae). Canad. Ent. 102: 513-5383.

RemMinctTon, C. L. 1958. Genetics of populations of Lepidoptera. Proc. Tenth Int.
Congr. Ent. 2: 787-805.

REMINGTON, C. L. 1968. Suture-zones of hybrid interaction between recently joined
biotas. Evol. Biol. 2: 321-428.

ScuppER, S. H. 1889. The butterflies of the eastern United States and Canada,
with special reference to New England. Vol. 1. Published by the author, Cam-
bridge, Mass., pp. 250-305.

Suaprro, A. M. anv J. D. Biccs. 1970. A hybrid Limenitis from New York. J.
Res. Lepid. 7: 149-152.

SreBBINS, G. L. 1958. The inviability, weakness and sterility of interspecific hy-
birds. Adv. Genet. 9: 147-215.

IDENTITY OF PHANETA REFUSANA (WALKER) WITH
DESCRIPTION OF A NEW SPECIES (TORTRICIDAE)

WILLIAM FE. MILLER

North Central Forest Experiment Station, USDA Forest Service,
St. Paul, Minnesota

The name Phaneta refusana (Walker) is currently used for moths
matching Heinrich’s (1923) idea of Walker's species. Although Heinrich’s
interpretation is the most explicit available, Heinrich never saw the Walker
type. He perpetuated Kearfott’s (1905a) identification which was based
mainly on Walsingham’s (1879) description and lithograph figure. Photo-
graphs of the holotype taken by N. S. Obraztsov at the British Museum
(Natural History), and made available by the American Museum of
Natural History, show that true refusana is actually different from the
refusana of Heinrich. I confirmed this finding by examining the type itself
at the British Museum. The misidentified moths have no valid name and
I here propose a new one for them.

The letter n in this paper signifies the number of specimens observed
for a particular statement. Values of n differ from the total number
studied because all specimens were not satisfactory for all purposes.
Forewing lengths (one wing) are given to the nearest 0.5 mm including
fringe and excluding patagium. The generic name Phaneta is used as
suggested by Obraztsov (1952).

Phaneta refusana (Walker), new combination

Grapholita refusana Walker, 1863.
Semasia refusana; Walsingham, 1879.

, a

i ee TI er I a rs

VoLUME 25, NuMBER 4 285

Figs. 1-3. Phaneta spp. 1, Phaneta refusana, wings of holotype male as photo-
graphed by Obraztsov (a) and Valva of holotype (b); 2, Phaneta verna, wings of
holotype male (a) and Valva (b); 3, Phaneta autumnana, wings of an example from
St. Clair Co., Mich (a) and Valva (b).

Walker and Walsingham mentioned one specimen (Fig. 1) which is a
holotype by reason of monotypy. The holotype, which is in the British
Museum, is labeled “Type H. T.; Hudson’s Bay St. Martin’s Falls G.
Barnston 1844-17; 44 17 St. Martins Falls; Grapholita refusana Wky.
neeebYPi ¢ deser,; 138. G...7....; B. M. ¢ Genitalia slide No. 4891.”
Its forewing measures 7.5 mm. True refusana is distinguished primarily
by its wide forewing and male valva which has only slight constriction in
width (Fig. 1). The species is known only from the type. I have located

286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

no examples in major North American collections nor in the British Mu-
seum. It may be a rare boreal species.

Phaneta verna Miller, new species

Thiodia refusana; Heinrich, 1923; Kearfott, 1905a; Kearfott, 1950b; McDunnough,
1939, no. 6782; McDunnough, 1942.

Head sordid white. Palpus white except for brown terminus and faint tinge of
orange on outer side. Antennal base, collar, and patagium sordid white, partly tinged
with yellowish orange. Thorax sordid white except first and second segments which
dorsally are light yellow. Legs sordid white, except fore and middle legs which are
tinged on outside with brownish yellow and have brown and white banded tarsi.
Forewing (Fig. 2a) 8.0 mm, predominantly light yellowish brown, the hue darkening
slightly on inner half and at base of fringe. Costa white except for about six faint
brown geminations. Outer half of forewing marked with thin silvery gray lines, one
of which partly encircles ocelloid area. Dorsal half of ocellus with 10 or 11 regularly
arranged black spots, ventral half sordid white due to white-tipped brown scales, and
surrounded by light yellow. Fringe speckled due to brown banded white scales. Hind
wing mostly white, grading to light brown at outer margin. Fringe mostly white with
base light brown and tip edged slightly with gray. Abdomen sordid white. Narrowest
width of valva about one-third the greatest width of cucullus.

The description is based on the holotype male which is in the American
Museum of Natural History. The holotype is labelled “Criddle Aweme
Man. 21V04; Kearfott Col. Ac. 4667; ¢ genitalia V.26.69 Slide 85 C. W.
Taylor.” The type locality is Aweme, Manitoba, Canada.

Besides the holotype, I studied 17 specimens from localities as follows:
MICHIGAN, Oakland, Allegan, and Ingham Counties; coLorapo, El Paso
Co.; NEw JERSEY, Middlesex Co.; CONNECTICUT, Windham Co.; ONTARIO,
Cochrane Co.; PENNSYLVANIA, Allegheny Co.; Nova scotia, Kings Co.
Forewings ranged from 7.0 to 8.5 mm (13 n).

Phaneta verna most resembles and is sympatric with P. autumnana
(McDunnough). It is tempting at first to think they are spring and fall
broods of the same species. This possibility must be ruled out by the
color and structural differences summarized below:

Item verna (7) autumnana (n)
Forewing Basal half light ( 8) Basal half dark ea)
Hind wing Light ( 8) Dark (@laiy)
Male genitalia Valva moderately con- Valva extremely con-

stricted in width stricted in width

(Fig. 2b) (il) (tie, So) (18)
Female genitalia Sternite laterally lobed Sternite not lobed near

near fusion with lamella fusion with lamella

antevaginalis ( 5) antevaginalis ( 3)
Flight period May 15-29 ( 7) Sept. 3-Oct. 13 (21)

(aN ie alas

(Michigan examples
only )
eaen yy]

VoLUME 25, NuMBER 4 287

Phaneta autumnana (McDunnough), n. comb.

Thiodia autumnana McDunnough, 1942.

This species is illustrated here for the first time (Fig. 3). I studied a
total of 25 examples from localities as follows: micHican, Osceola, Ma-
comb, Livingston, St. Clair, Otsego, Midland, and Shiawassee Counties;
WISCONSIN, Oneida Co.; CoNNEcTicuT, Windham Co. Forewings ranged
from 7.0 to 8.5 mm (24 n).

Acknowledgment

I thank the following for help with this study: Frederick H. Rindge,

American Museum of Natural History; Thomas N. Freeman, Entomology

Research Institute (Canada); Jerry A. Powell, University of California;
and Paul E. S. Whalley, British Museum (Natural History).

Literature Cited

HernricuH, C. 1923. Revision of the North American moths of the subfamily
Eucosminae of the family Olethreutidae. U. S. Nat. Mus. Bull. 123.

KearFotr, W. D. 1905a. Assiniboia Micro-Lepidoptera, collected by Mr. T. N.

Willing. Can. Ent. 37: 41-48, 89-93.

1905b. Manitoba Micro-Lepidoptera. Can. Ent. 37: 205-209, 253-256,
293-296.

McDunnoucu, J. 1939. Check list of the Lepidoptera of Canada and the United
States of America. Part II. Microlepidoptera. So. Calif. Acad. Sci. Mem. 2.

1942. Tortricid notes and descriptions. Can. Ent. 74: 63-71.

Osraztsov, N. 1952. Thiodia Hb. as not a North American genus (Lepidoptera,
Tortricidae). Ent. News 63: 145-149.

WALKER, F. 1863. List of the specimens of lepidopterous insects in the collection
of the British Museum. Part 28. Tortricites and Tineites.

WALSINGHAM, T. DE GREY, SIXTH LORD. 1879. Illustrations of typical specimens of
Lepidoptera Heterocera in the collection of the British Museum. Part 4. North-
American Tortricidae.

THE LIFE HISTORY OF HELIOLONCHE PICTIPENNIS
(NOCTUIDAE)

D. F. Harpwick

Entomology Research Institute, Canada Department of Agriculture,
Ottawa, Ontario

Heliolonche pictipennis (Grote, 1875, p. 220) feeds in the larval stage
on the Desert Dandelion, Malacothrix glabrata (A. Gray) (Fig. 2). In
the spring of the year when its food plant is in blossom, the moth often

288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

becomes locally very abundant in some areas of the dry interior of southern
California. All the specimens in the Canadian National Collection were
taken on the southern California deserts between the middle of March
and the middle of May, except for a single specimen taken at Tucson,
Arizona, on August 10.

Malacothrix glabrata is an annual composite with yellow blossoms which
is recorded (Munz, 1963) as being distributed from southern California
northward to Idaho and eastward to Arizona. Whether the Desert Dan-
delion extends eastward to the Tucson area and if so whether it germinates
in response to summer rains in that area is not known.

In the spring of 1955 near Victorville, California, a pair of Heliolonche
pictipennis was found in copula in the blossom of another annual yellow
composite, Glyptopleura setosula Gray. It was not determined whether
this species constitutes an alternative host plant or whether the association
of the moth and the blossom was purely a fortuitous one.

Behaviour

Heliolonche pictipennis is an exclusively diurnal species and in desert
areas in which its food plant is abundant and in early blossom, the little
moth may usually be found without difficulty flying swiftly from blossom
to blossom or resting in the flowering head. When resting or copulating
on the head, the moth is usually difficult to discern because it closely
resembles the red “button” in the centre of the Malacothrix blossom. In
the late afternoon the ray petals of Malacothrix close upward and inward
over the middle of the head and moths that have come to rest on the
blossom are thus enclosed until the following day.

The oviposition pattern of pictipennis is similar to that of Heliolonche
carolus described by Hardwick (1969), and the eggs are inserted between
the florets from the upper surface of the head. The ovipositing female
does not achieve the depth of penetration of the female carolus, however,
and usually the eggs come to rest among the bristles of the pappus well
above the developing seeds.

Four wild-caught females of pictipennis deposited a mean of 28.5 eggs,
and the maximum laid by a single female was 41. The majority of eggs
observed hatched on the fifth day after deposition.

The larva feeds at first on the contents of the florets and subsequently
attacks the seeds. During one of the median stadia, the larva usually
quits the first head and enters a second in which it completes its develop-
ment. As with other species of Heliothidinae, the mature larva enters the
ground to pupate and it is in the pupal stage that the species spends the
greater part of the year.

VoLUME 25, NuMBER 4 289

: — . “J re aS =

Figs. 1-4. Heliolonche pictipennis (Grote) and plants with which it is associated.
1, Adult, Adelanto, Calif.; 2, its food plant, Malacothrix glabrata (A. Gray); 3, pupae;
A, Glyptopleura setosula Gray in which a copulating pair of pictipennis was found.

Descriptions of Stages

The following descriptions of immature stages were based on the prog-
eny of four females taken in the White Water Pass area north of Palm
Springs, California. The larvae were reared individually at room tem-
perature on the flowers and seeds of Malacothrix glabrata. Rearing tech-
niques employed were those outlined by Hardwick (1958). The estimate
of variability following the means for various values is the standard
deviation.

Adult (Fig. 1). Vestiture of head and thorax olivaceous fawn or fawn-grey.

Abdomen dark brown with a fawn anal tuft and often with fawn segmental rings.
Underside of thorax and abdomen dark brown with varying amounts of fawn over-

290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

scaling. Forewing dull yellow and light fawn marked with pink. Transverse anterior
line smooth, weakly excurved. Basal space fawn, variably suffused with pink, most
strongly so along inner margin of t.a. line. Transverse posterior line shallowly sinuate,
weakly excurved around cell, then usually weakly incurved to trailing margin. Median
space dull yellow with a narrow, fawn or pink, costal band. Orbicular spot absent.
Reniform spot usually evident as a narrow dark shade. Subterminal space narrow,
usually suffused with pink. Terminal space pale olivaceous fawn. Fringe light brown,
variably suffused with pink. Hind wing black, variably marked with white; white
occupying entire central area of wing or reduced to a narrow evanescent median band.
Fringe white with a brown basal line. Underside of both wings pale grey marked
with brown. Forewing with a brown basal dash, narrow reniform spot, and incom-
plete subterminal band. Hind wing with a brown basal patch, anal patch, and inner
marginal band.

A very pale form of pictipennis also moderately common in which pink absent, fawn
coloring replaced by silvery-grey, and yellow replaced by white.

Expanse: 15.3 + 1.2 mm (78 specimens ).

Egg. Very pale yellow when deposited. Turning somewhat darker yellow on day
after deposition, then remaining essentially unchanged until a few hours before
hatching when mandibles and then ocelli becoming visible through chorion.

Dimensions of egg: length, 0.919+ 0.034 mm; diameter, 0.519 + 0.040 mm
(10 eggs).

Incubation period: 4.8 + 0.4 days (60 eggs).

First-Stadium Larva. Head orange-brown, variably suffused with chocolate-
brown. Prothoracic shield fawn, usually heavily suffused with smoky-brown. Suranal
shield dark smoky-brown. Trunk pale cream. Thoracic legs smoky-brown. Spiracles
with dark-brown rims.

Head width: 0.262 + 0.012 mm (20 larvae).

Duration of stadium: 3.3 + 0.7 days (29 larvae).

Second-Stadium Larva. Head smoky-brown. Prothoracic and suranal shields
concolorous with head. Trunk greyish-cream, becoming toned with yellow after larva
resumes feeding. A dark mid-dorsal band and paler subdorsal areas usually evident.
Spiracles with dark-brown rims. Thoracic legs smoky-brown.

Head width: 0.443 + 0.022 mm (22 larvae).

Duration of stadium: 2.4 + 0.7 days (29 larvae).

Third-Stadium Larva. Head orange-brown, heavily suffused and mottled with
dark brown; mottling often so heavy as to almost obscure lighter colouring. Pro-
thoracic shield dark brown with a cream median line, and usually with broader, cream
submarginal lines. Suranal shield light orange-brown, variably suffused and mottled
with dark brown; usually a pair of cream submarginal lines evident. Mid-dorsal band
of trunk varying from light chocolate-brown to orange-brown, and usually with a dis-
continuous, cream median line. Subdorsal area white or cream with a median orange-
brown band; median band paler than mid-dorsal band. Supraspiracular area con-
colorous with mid-dorsal band, and with a discontinuous, white or cream, median
line. Spiracular band white or pale cream with a discontinuous, orange-brown median
line. Suprapodal area greyish-fawn, irregularly marked with light orange-brown. Mid-
ventral area fawn-grey. Spiracles with dark-brown rims. Thoracic legs dark greyish-
brown.

Head width: 0.715 + 0.024 mm (25 larvae).

Duration of stadium: 2.4+0.5 days (29 larvae).

Fourth-Stadium Larva. Head cream or pale fawn, mottled with orange-brown,
and with a few black spots. Prothoracic and suranal shields light orange-brown,

ariably marked with black; prothoracic shield with three longitudinal white lines,
and suranal shield with two longitudinal white lines. Mid-dorsal band of fk
medium chocolate-brown with a pale median shade. Subdorsal area white or cream

ha a of longitudinal lines of orange or light orange-brown. Supraspiracular area
to dark brown, often darker than mid-dorsal band; with an irregular and

VoLUME 25, NUMBER 4 291

Figs. 5-8. Heliolonche pictipennis (Grote), fifth-stadium larvae. 5, 6, Dorsal:
7, 8, left lateral.

discontinuous, pale median line. Spiracular band white with an irregular and dis-
continuous, orange-brown median line. Suprapodal area varying from orange-brown
to medium chocolate-brown, with white arcuate markings. Mid-ventral area yellowish-
grey or brownish-grey. Spiracles with dark-brown rims. Thoracic legs pale fawn or
pale grey, variably suffused with dark brown.

Head width: 1.14 + 0.03 mm (25 larvae).

Duration of stadium: 3.8 + 0.9 days (29 larvae).

Fifth-Stadium Larva (Figs. 5-8). Head pale fawn or cream, mottled dorsally
with chocolate-brown and with several black spots on face. Prothoracic shield fawn
suffused with black; in some specimens black suffusion so heavy as to obscure fawn
colouring; shield with two or three white longitudinal lines. Suranal shield poorly
distinguished from remainder of trunk. Maculation of trunk complex. Mid-dorsal
band pale pink, emarginated laterally by irregular lines of brown or red. Subdorsal
area white or pale cream with a pair of pale-red, median longitudinal lines. Supra-
spiracular area brown with an irregular white median line or shade; spiracular band
often becoming grey toward posterior margin of each segment and thus with a patchy
appearance. Spiracular band broad, white, with a segmentally interrupted, pink or
light-red, median line. Suprapodal area fawn-grey, demarked from spiracular band
by a light-red line; suprapodal area with an irregular pattern of white and red arcuate
marks. Mid-ventral area grey. Spiracles with dark-brown rims. Thoracic legs cream
or fawn, lightly marked with chocolate-brown.

Head width: 1.84 + 0.06 mm (7 larvae).

Duration of feeding phase of fifth stadium: 4.8 + 1.1 days (29 larvae).

Duration of prepupal phase of fifth stadium: 4.5 + 0.9 days (15 larvae).

Pupa (Figs. 3, 9, 10). Light brown with a strong green suffusion on head and
thoracic appendages. Spiracles on abdominal segments 5, 6 and 7 borne in shallow
depressions of cuticle, remainder on a level with general surface of cuticle. Spiracular

292. JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Figs. 9, 10. Heliolonche pictipennis (Grote), apical abdominal segments of pupa.
9, Ventral; 10, right lateral.

sclerites narrow. Anterior marginal areas of abdominal segments 5, 6 and 7 sparsely
and shallowly pitted. Proboscis terminating a short distance anterior to apexes of
wings. Cremaster consisting of four slender, elongate, well-spaced setae borne in a
single row at narrowly rounded apex of tenth abdominal segment.

Length from anterior end to posterior margin of fourth abdominal segment: 6.4 +
0.3 mm (19 pupae).

Acknowledgments

I am grateful to Mr. John E. H. Martin of this Institute for the photo-
graphs accompanying this paper, and to my associate, Mr. Eric Rockburne,
for measuring the immature stages and drawing pupal structures.

Literature Cited

Grote, A. R. 1875. Supplement to the list of North American Noctuidae. Bull.
Buffalo Soc. Nat. Sci. 2: 209-223.

Harpwicx, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed
species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae.
Can. Ent. Suppl. 6.

1969. The life history of Heliolonche carolus (Noctuidae). Jour. Lepid.
Soc. 23: 26-30.
Munz, P. A. 1963. A California flora. University of California Press, Berkeley.

VoLuME 25, NuMBER 4 293

AGUNA CLAXON (HESPERIIDAE) NEW TO THE UNITED STATES

On October 21, 1970, while collecting in the Santa Ana Wildlife Refuge, Hidalgo
Co., Texas, I took a single specimen of Aguna claxon Evans, a large skipper with green
gloss above and with a white band across the secondaries below.

Evans (1952, Cat. Amer. Hesp. B. M.) separated A. claxon from A. coelus (Stoll).
A. coelus has the hind wings short-tailed and is less vividly green above. A. claxon
has the hind wings lobed rather than tailed, and is more brilliantly green above. There
are also differences in the male genitalia. According to Evans, A. coelus does not
occur in Mexico, but ranges from Central America far into South America. A. claxon
occurs in Mexico and south into Central America, where both species occur.

In Godman & Salvin (1893, Biol. Centr.-Amer. Lep.-Rhop. 2: 287), A. claxon ap-
pears as Goniurus coelus (Cramer). In Hoffman’s List (1941, An. Inst. Biol. 12(1):
244) A. claxon is listed as A. coelus (Cramer). According to Evans, the figures in
Seitz, Amer. Rhop., are mixed. Of the figures referred to as Goniurus caelus (sic!)
Cramer, the underside is stated to be correct. The upper side is that of Aguna aurunce
( Hewitson ).

The specimen of Aguna claxon was taken in shade, at the flowers of Eupatorium
odoratum L.., in the heat of the day. On the same plant, a single specimen of Bolla
brennus (Godman & Salvin) was also taken.

J. W. TitpEn, 125 Cedar Lane, San Jose, Calif.

REMARKS ON “THE USE OF NET-TRAPS AT PALAWAN, PHILIPPINES”

Mr. Jumalon’s note under the above heading (1970, Journ. Lep. Soc. 24: 303-4)
brings out some interesting contrasts between results there and in East Africa.

In East Africa, carnivore dung, that of lion, leopard, civet, etc., is well known for its
attraction to many male nymphalids, and man, after all, is mainly a carnivore. It is
perhaps worth recording here that the droppings of the domestic dog, although fed
largely on meat, do not seem to be attractive.

The usual fruits used as bait in East Africa are banana, pineapple and mango;
papaya is mentioned in literature, but I have never found it nearly as attractive as
the first three. I have never heard of Custard Apple (Annona squamosa) being used
as bait.

The trapping of Papilionidae, Pieridae and Hesperiidae is entirely contrary to my
experience in East Africa and I am inclined to think that these were cases of low-flying
butterflies getting under the edge of the very large net and failing to get out again, a
situation analogous, perhaps, to the trapping of birds in mist-nets, rather than attraction
to bait.

In East Africa the only species attracted to fruit baits belong to the Satyridae,
Nymphalidae (mainly Charaxinae, Nymphalinae and Eurytelinae, with Neptidinae,
Vanessinae and Argynnidinae to a lesser extent) and Libytheidae, and males of these
species are attracted to faeces.

Almost all families are attracted to damp mud, but again males only. I have very
occasionally found female nymphalids on patches of mud but am of the opinion that
there is a primary attraction in the form of fermented fruit juices in such cases. The
late C. L. Collenette was of the opinion that the major attraction of damp mud was its
salt content, and pointed out that the most attractive areas were the banks of streams
where clothes were washed or where humans and animals forded streams. Certainly,
in my experience, the attraction of damp mud increases with the distance from the
sea, it is far more attractive in Uganda than it is on the Kenya coast.

D. G. SEvAstorpuLo, Mombasa, Kenya.

294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

NOTES ON UNUSUAL SPECIES OF LEPIDOPTERA
FROM SOUTHERN FLORIDA

Danaus eresimus Cramer. One worn virgin female was taken while feeding at
flowers of Asclepias curassavica L. on June 6, 1970 in South Miami, Dade County.
Another specimen, a male in nearly fresh condition, was taken at flowers of Eupatorium
serotinum Michx. on October 6, 1970 near Homestead Air Base in southern Dade
County. Both individuals apparently belong to populations of D. eresimus tethys
Forbes which is common in the greater Antilles.

A search of leaves and flowers of Asclepias curassavica, A. tuberosa rolfsii ( Britt.)
Shiners, and Sarcostemma clausa Vail in the vicinity of the capture sites and elsewhere
in Dade County yielded only larvae and eggs of resident Danaus gilippus berenice
(Cramer) and D. plexippus L. If D. eresimus is established in Florida, it is rare. It
is more likely that individuals stray in from the West Indies as suggested by Kimball
(1965, Lepidoptera of Florida).

Urbanus dorantes Stoll. This species has recently been reported from southern
Florida (Clench, 1970, J. Lepid. Soc. 24: 240-244). U. dorantes is well established
in Dade County and flies at least between July and April, encompassing three or more
broods. From November to January of 1969 and 1970, U. dorantes and U. proteus L.
could be taken with equal frequency in Coral Gables, South Miami, and in sawgrass
marshes near Pinecrest, Monroe County. Females of U. dorantes oviposit on Des-
modium tortuosum (Sw.) DC as do U. proteus females.

Tuomas E. Puiske, Department of Biology, University of Miami, Coral Gables,
Florida.

DANAUS PLEXIPPUS (NYMPHALIDAE) ATTACKING
RED-WINGED BLACKBIRD

Last summer, Mr. James Erickson and I were collecting female monarchs, Danaus
plexippus L., in order to establish a laboratory culture for our study of larval feeding
efficiencies. On July 6 and 7, 1970, we were collecting in a large (approximately 10
acres ), rather pure stand of milkweed, Asclepias syriaca L., at the Ithaca, New York
old airport (Tompkins Co.). In one corner of the stand we were constanly harassed
by a male red-winged blackbird, Agelaius phoeniceus L., on whose nesting territory
we were apparently trespassing. In three separate instances we saw a male monarch
“attack” the red-winged blackbird as it hovered over us at a height of some 20-30
feet. Flying and diving at the bird for less than a minute each time, the monarch
appeared to “frighten” the bird, but although backing off slightly, the bird never left
the area. The monarch would break off the encounter and come swooping down to
the milkweed patch, flying rather rapidly.

Clark (1931, Butterflies of the District of Columbia) mentioned that male monarchs
have been seen to attack smaller butterflies and small birds (i.e., hummingbirds and
warblers). The red-winged blackbird eats both seeds and insects, especially beetles,
grasshoppers, and caterpillars. The monarchs may have been “flaunting” their
warning coloration, although if one assumes that they had fed as larvae on these milk-
weed plants, they would be palatable to predators because A. syriaca lacks cardiac
glycosides (Brower, 1969, Scientific American 220: 22-30).

FRANK SLANSKy JR., Department of Entomology and Limnology, Cornell Univer-
sity, Ithaca, New York.

VoLuME 25, NuMBER 4 295

A SIMPLE METHOD FOR PREPARING MALE HESPERIID GENITALIA
FOR EXAMINATION WITHOUT DISSECTION

Examination of the genitalia of Lepidoptera usually involves dissection, a rather
unesthetic treatment for a mounted specimen. Evans, in preparing his catalogues of
the Hesperiidae, frequently used a dry dissection method, but this is not always totally
satisfactory. For the past year I have been extruding the male genital armature while
specimens are still fresh by gently pulling on the clasps until the entire armature
“pops out.” This has not always proved satisfactory because as the specimen dries, the
genitalia frequently retract slowly, and in any event, the clasps remain at least partially
closed necessitating some tissue rupture to open them for examination of the inner
faces and the penis, uncus and gnathos.

Further experimentation has indicated that the clasps can be held in a wide open
position during drying by applying a bit of Duco or similar cement, which can be re-
moved later. The technique is as follows:

Holding the fresh insect by the thorax with forceps in normal pinching position,
the genital armature can be extruded by gently pulling out the clasps with fine for-
ceps. Sometimes the uncus will be bent down covering and distorting the gnathos;
this usually can be teased into a normal position with a dissecting needle. Once the
genitalia are fully exposed, grasp the abdomen with fine curved forceps immediately
forward of the vinculum which further spreads the clasps. A small quantity of cement
is then smeared over the area of the junction of the clasps which are held spread wide
with a second pair of fine forceps until the cement dries, 5 minutes or less. I prefer
to spread the clasps to a position normal to the abdomen as it simplifies photography.
When the specimen is thoroughly dried, either spread or in papers, the cement is
easily removed; usually it can simply be peeled away. However, if stuck too firmly,
it can be dissolved away by washing in acetone, leaving the genitalia well exposed
for study.

The same process also can be applied to relaxed specimens provided the genitalia
have been previously extruded, but is not as successful as with fresh specimens.
Because of the general structure of the male genitalia of the Hesperiidae, the method
is especially applicable to this family though of limited use in others, where for
example, details of the anterior portions of the tegumen, the base of the penis or the
structure of the saccus are important features. In addition, in several families of butter-
flies, the genitalia are either too lightly sclerotized or too difficult to extrude to lend
themselves to this process.

I sincerely hope that others may find this method as useful as it has been to me.

STEPHEN R. STEINHAUSER, Apartado 109, San Salvador, El Salvador

CAPTURES OF ERORA LAETA IN NORTH CAROLINA (LYCAENIDAE)

Roever (1962, J. Lepid. Soc. 16: 1-4) described several records for Erora laeta
(Edwards) from the southern states. These included a single capture in Tennessee
(April 15) and a single capture in North Carolina (July 17). Clark and Clark (1951,
Butterflies of Virginia) list a single capture in Virginia (June 23) and since that pub-
lication at least two additional spring specimens have been taken. There are no re-
corded specimens from Georgia or northeastern Alabama. All Erora laeta taken in the
southern states were found in the mountain regions.

On July 1, 1970, I drove from Durham, North Carolina, to Alleghany County in the
northwestern corner of the state. I had been in the area a week earlier but rain had
cut short my collecting. At that time the Speyeria were in good flight and I had made
the trip in hopes of getting a nice series of cybele, aphrodite and especially idalia. I

296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

arrived at my favorite collecting spot (elevation 2700 feet, County Road 1345) about
noon but soon discovered that the clover field which had been alive with Speyeria the
previous week was now very dead. So, I continued down the road in order to check
several other spots that had been productive in the past. One such place was a small
patch of Ceanothus americanus bordering the road. This time I decided to follow the
Ceanothus over a barbed wire fence and up a steep slope. A few specimens of Strymon
falacer and S. titus mopsus were taken and Speyeria females were scattered through
the area. Much to my surprise the Ceanothus actually covered an acre or more but
only a small patch was visible from the road. I worked my way up and down the slope
adding a few Strymon and Speyeria each trip. In addition, two Strymon liparops were
taken. On one of these trips my vision happened to fall right on an Erora laeta sipping
nectar in the middle of a large patch of Ceanothus! The slope had scattered trees and
shrubs on it, but this was the most open part. After staring in disbelief for several
seconds (knowing of course that it would disappear forever), I came to my senses
and netted the specimen. It was a fresh female! I searched the area thoroughly for
the next hour but found no additional specimens. Beech, the supposed foodplant
(but why? ), was not located near the Ceanothus nor in the immediate area. I decided
to drive to a location in Ashe County where S. idalia was often common. This location
(on U. S. 221 near the junction with County Road 1570) is also on a hillside at 2700
feet elevation, but Ceanothus is scarce. Instead there is a good colony of Asclepias
tuberosa and the Speyeria were busily flying from one plant to the next. I joined the
merriment taking idalia and a number of somewhat worn aphrodite and cybele.
Actually S. aphrodite was the most common fritillary and a half dozen could be taken
off a single flowerhead. At one such clump of orange milkweed I patiently waited
for a number of aphrodite to settle so that I could maximize my effort. Just when 5 or
6 would settle down, an idalia would charge the group and mayhem would result.
Finally, in frustration I swung just as an idalia was approaching. I quietly cursed at
seeing only three aphrodite in the net. But, unbelievably there was an Erora laeta in
with them! Apparently it had been nestled among the aphrodite on the flowerhead,
and I had taken it unknowingly! This one was a fresh male. Additional searching of
the area produced no additional Erora nor were beech trees located. Both specimens
had been taken on flowers in open areas with scattered trees and shrubs, but in full
sunlight. The species is reported to be a denizen of beech forests.

These captures, some 13 air miles apart, may only reflect an extraordinary amount
of luck. Smith (1960, J. Lepid. Soc. 14: 239-240) took 7 laeta in New Hampshire
in an area where they had not been found previously. Perhaps both encounters rep-
resent local population “pops” (with laeta you can’t call it an explosion! ). Clark and
Clark (1951) suggested there might be three broods in Virginia. This seems like a
good possibility, but I do not know of any August or September records. Until we
know more about the life-history and habits of Erora laeta, it will remain a rare species.
However, it must be sufficiently common to maintain a breeding population, and I
suspect that finding it is only a matter of knowing where to look. Of course having
captured two in one day, 13 miles apart, and one by accident, I have given up hope
of ever finding another!

J. Botxinc Sutuivan, Duke Marine Laboratory, Beaufort, North Carolina.

in i

~ i

ee

VoLUME 25, NUMBER 4 2 3)7f

BOOK REVIEW

BUTTERFLIES OF TRINIDAD AND Tosaco. Malcolm Barcant. Collins Publ., London.
1970. 314 pp. + 28 plates. Approximately $6.00.

In the preface, Mr. Barcant states that his book “is prepared in a somewhat
new ... style’ that he believes will become popular. I heartily agree that it is a new
style, for what other book has divided its subjects by such ambiguous categories as
“butterflies of the home garden,” “fruit and sap suckers,” “migrants,” “locals and
brooders,” “semi-rare species,” “species becoming rarer,” or “the remaining Lycaenids”?
Add to this the fact that there are no keys or alphabetical indexes, and you can imagine
the problems of using the text as a field guide. A particular butterfly may be si-
multaneously a “butterfly of sunshine and flowers” (chapter 6F ), “mud-puddle butter-
fly” (chapter 6D), “migrant” (chapter 6F), “southern resident” (chapter 61), “well
known species” (chapter 9A), “highly prized species” (chapter 9D), “species becoming
more common” (chapter 9E), “species with special interests” (chapter 9G), and a
“remaining Lycaenid” (chapter 10), but will be discussed under but one of these
categories. For Trinidad butterflies, the situation is complicated even further by nu-
merous groups of very similarly marked species. Papilio thoas, for instance, is treated
as a home garden butterfly” (page 74) with no cross reference to the almost identical
Papilio homothoas (a species “becoming rarer” on page 188) or to the similarly marked
Papilios androgeus and lycophron (“water drinkers” on page 92). For somebody un-
familiar with Neotropic butterflies, the only way to identify a specimen with complete
assurance would be to read the entire book. I do not share Mr. Barcant’s hope that
this style of book will become more popular.

There is, however, quite a bit of excellent information in the book. Mr. Barcant has
collected on Trinidad for many years and his comments on the habitat and habits of
each species should be of considerable value to the student of Neotropical Lepidoptera,
as should the citations of larval foodplants. One chapter deals with “flowers popular
among adult nectar-sucking butterflies,” which, aside from the practical collecting
aspects, is valuable information to have recorded. Another section deals with collecting
localities and seasons on Trinidad and would be of considerable value to somebody
planning a collecting expedition to the island. Nearly all of the Trinidad species,
excluding Hesperiidae, are satisfactorily reproduced in color plates, with however, a
reduction in size and no scale or other indication of actual size cited.

Aside from its general deficiency as a field guide, the book has a number of other
weaknesses. Typographic errors, especially in the latin names of species, are frequent,
e.g. Anteos clorinde is consistently cited as “chlorinde’, Papilio lycophron as “ly-
crophon,” Heliconius hecale as “hecali,’ etc., and it would be impractical to list all
of these. There are also a number of erroneous statements such as “each species of
butterfly is constant and its sex organs differ in shape from any other species so that
mating outside its own female is not possible” or “it is necessary to add brackets to
the name of the man who first described the butterfly.”

I also believe that Mr. Barcant exercised poor judgement in introducing several spe-
cies as new. Pachthone barcanti, which was described by G. E. Tite in 1968, is in-
troduced as Sp. Nov. with no author cited. Adelpha cytherea insularis and Sostrata
pusilla manzanilla are also introduced as Sp. Nov. with no author citation. Unless
quite recently, these have not yet been published elsewhere and, if this is so, the
descriptions in the text constitute original description. I am sure that this was not
intended to be the case and I doubt if Barcant was designed to be their author.

A check-list of Trinidad butterflies (chapter 14) is arranged along no phylogenetic
order in current use as follows: Satyridae, Danaidae (including Acraea), Ithomiidae,
Heliconidae, Nymphalidae, Morphidae, Brassolidae, Libytheidae, Papilionidae, Pier-
idae, Riodinidae, Lycaenidae and Hesperiidae. The generic names used are sadly
dated for the Lycaenidae and Hesperiidae, but pretty much in current usage for other
families. In the Nymphaloid groups the only readily apparent errors, in this respect,

298 JOURNAL OF THE LEPIDOPTERISTS SOCIETY

are the use of Callicore for Diaethria and, in turn, the retention of Catagramma for
Callicore.

English vernacular names, such as Flambeaus, Pages, Shoemakers, Crackers, Cattle-
Hearts and others, that are in actual use in Trinidad are recorded, perhaps for the first
time. In spite of their lack of scientific value, I find them quite interesting and am
glad to see them preserved by publication.

There is much of interest and value in the book; the serious student of Neotropic
Lepidoptera may find it a valuable reference. However, it was intended primarily as
a field guide and aid for the young collector on Trinidad; a usage for which it has
some very serious limitations.

Joun H. Masters, Lemon Street North, North Hudson, Wisconsin.

The most serious fault with the book is one that cast a question over all the book
contains. There are three names used in the book that are labeled “Sp. Nov.” One of
these is Pachythone barcanti which was described by G. E. Tite of the British Museum
in 1968 and certainly Barcant knew that this was not a nameless species to which he
was giving a name in this publication. The other two “Sp. Nov.” are equally senseless.
Adelpha cytherea insularis was described by Fruhstorfer in “1915” 1916 in Seitz’
Macrolepidoptera, 5: 521. Barcant was familiar with this book and used it. Sostrata
pusilla manzanilla was described by Kaye in 1940 and certainly anyone writing on
Trinidadian butterflies must have examined all of Kaye’s writings about them. I don't
know whether these actions of declaring named species “Sp. Nov.” was done in ig-
norance or with the supposition that no one would bother to investigate.

F. M. Brown, Fountain Valley School, Colorado Springs, Colo.

BOOK REVIEW

THE Motus or AMERICA NortH OF MExico, FAscICLE 21, SPHINGOIDEA, by Ronald
W. Hodges. 1971; 158 pp. + i-xii, 14 coloured plates; paper-bound. E. W. Classey
Limited and R.B.D. Publications Inc. Distributed in North America by Entomological
Reprint Specialists, P.O. Box 77971, Dockweiler Station, Los Angeles, California.
Price $24.00 U.S. (Subscription Price $19.60).

This is the first published fascicle of the eagerly awaited series of the North Amer-
ican Heterocera. Dr. Hodges is to be congratulated on his very fine treatment of the
115 species of hawk moths in the boreal American fauna. Under each species heading
is given a brief synonymical bibliography, a description of diagnostic structural and
macular features, a listing of larval food plants and a discussion of the distribution.
Keys to genera and species are presented, and these in most cases seem easy to use.
Old keys to genera based on pupae (Mosher, 1918) and larvae (Forbes, 1911) are
reproduced in an introductory section. The nomenclature of the various structural
characters employed in classification is well explained both in the text and by line cut
illustrations. The most outstandingly laudable feature of the volume, however, are the
magnificent coloured plates; the specimens are all ready to fly right out of the pages.

This is a book that should be readily available to both the professional and the
serious amateur.

Pan)

D. F. Harpwicx, Editor.

VoLUME 25, NUMBER 4 299

A FreELD GUIDE TO THE BUTTERFLIES AND BURNETs OF SPAIN, by W. B. L. Manley
and H. G. Alleard. 1970. 182 pp., 41 colour plates including frontispiece. Pub-
lishers: E. W. Classey Ltd., Middlesex, England. (Available in U. S. from Entomo-
logical Reprint Specialists, P. O. Box 77971, Dockweiler Station, Les Angeles, Cali-
fornia 90007; price $37.50. )

This book includes all 229 species of butterflies known to occur in Spain, and in
addition to these, the 26 species of bumets (Zygaenidae). The colourful burnets
being diurnal are as popular with collectors as many of the butterflies and their in-
clusion enhances the usefulness of the book.

The term “field guide” usually carries with it the connotation of a compact volume
with detailed and comparative descriptions of the various species but neither of these
qualities is found in the “Butterflies and Burnets of Spain.” All species are illustrated
in beautifully prepared colour plates. In many cases, males, females, undersides, and
pattern variations are all reproduced at life size. Unfortunately the page number
for the text of the species is not included in the plates and it is necessary to refer to
the index to find the text reference for the butterfly. The plates do however include
the complete data for each specimen illustrated.

The text for each species is short and usually covers three subjects. These are: the
localities where the species may be most easily located, complete with elevation and
time of year, secondly, a discussion of the various subspecies and forms described,
and thirdly, the larval host plants.

In addition to the butterflies and burnets of Spain, sections at the end of the text
deal with the Balearic Islands, the Canary Islands, and Madeira with a similar text
and plates.

At the end of the text there is a complete systematic check list of species, subspecies,
and forms in the Iberian Peninsula, the Balearic Islands, the Canary Islands and
Madeira. The list includes references to original descriptions for a more detailed study
of the species.

“The Butterflies and Burnets of Spain” will be an invaluable field guide to collecting
as well as identification for the lepidopterist interested in collecting in any of the
areas treated in the book.

J. DonaLp LAFONTAINE, 916 Innswood Drive, Ottawa, Ontario.

300

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

INDEX TO VOLUME 25
Subject Index

behaviour, 2, 6, 20, 53, 80, 84, 109, 126,
AA NAG lS ORmiM pmol oan ae one
294.

book reviews, 86, 148, 152, 176, 221, 297,
298, 299

collections, 82, 83, 143, 168

Comstock, J. A., 25

distribution, 19, 29, 80, 84, 87, 114, 139,
143) TA. 149) a0 AS52220) 2465256"
293, 294, 295

habitats, 216

host plants, 6, 64, 146

hybridization, 68, 278

Janse; vA sa ie el

larvae, 262, 266

life histories, 1, 53, 58, 109, 126, 177, 181,
262, 274, 287

light trapping, 150

migration, 124

neutron irradiation, 238

new record, 293

nomenclature, 142, 264

obituaries, 211, 215

population structure, 22

presidential address, 155

rearing, 6, 68, 143, 247

regional lists, 29, 73, 137, 139, 150, 213,
222,

techniques, 22, 65, 83, 239, 293, 295

variation, 108, 143, 185, 234, 266, 271

Name Index

(New names in boldface )

Acrocercops, 198
agrifoliella, 201
Aguna, 293
alexandrae, 58
amanda, 53
Amatidae, 42
anicia, 246
Anisota, 84
anthedon, 256
Antheraea, 238
antiochella, 204
Apaturinae, 140
aphrodite, 150
Arcas, 87
Archichlora, 171
archippus, 278
Argynninae, 141
Arctiidae, 42
aristodemus, 126
arthemis, 278
astyanax, 278
Automeris, 234
bahamensis, 188
bohartiella, 200
Boloria, 84, 149
borealis, 256
Caloptilia, 201
Cameraria, 209
cardui, 147
castalia, 124

Cercyeuptychia, 13
Cercyonis, 12
Charaxinae, 140
Citheroniidae, 84
claxon, 293
Colias, 108
Coptodisca, 194
creola, 145

cupes, 109
Danaidae, 41, 214
Danaus, 294
delphia, 101
demodocus, 271
demylus, 149
deserticola, 109
dorantes, 294
dospassosi, 190
eleuchea, 185
emarginana, 115
Epinotia, 115
eresimus, 294
Erora, 295
eucalypti, 238
Euchloe, 64
eunomia, 84, 149
Euphydryas, 246
Euristrymon, 80
gemellus, 149
Geometridae, 169, 262, 266
Geometrinae, 169

VOLUME 25, NUMBER 4

Glaucopsyche, 240

Gracilariidae, 194

Heliconiinae, 141

Heliolonche, 287

Heliothis, 1, 264

Heliozelidae, 194

Hesperiidae, 20, 34, 213, 293, 295

Hyalophora, 68

insulariella, 198

io, 234

Itame, 266

Ithomiidae, 149

jacksoni, 171

jivaro, 105

jutta, 150

kershawi, 114

Kricogonia, 124

laddi, 84

laeta, 295

Lasiocampidae, 43

Lethe, 145, 256

Libytheidae, 38, 214

ligeae, 274

Limenitidinae, 140

Limenitis, 146, 278

Lithocolletis, 204

Lycaenidae, 37, 80, 87, 214, 240, 295

Lygris, 262

Lymantriidae, 53

Marpesia, 185

Megalopygidae, 43

Megathymidae, 34

Melitaeinae, 141

Mestra, 146

Morpho, 223

Neurobathra, 200

Noctuidae, 42, 109, 177, 181, 264, 274,
286

nokomis, 44

Notodontidae, 43

Nymphalidae, 19, 39, 44, 84, 114, 139,
143, 146, 149, 185, 214, 246, 247, 278,
294

Nymphalinae, 140

Ocnerogyia, 53

Oeneis, 150)

301

olympia, 64

ontario, 80
Ornithoptera, 58
Papilio, 1265 1427 271
Papilionidae, 36, 73, 126, 142, 214, 271
Patricia, 149

Phaneta, 284

philodice, 108
Phyciodes, 143

piasus, 240

pictipennis, 287
Pieridae, 6, 36, 64, 108, 124, 144, 214
Pieris, 144

Plebejinae, 240
plexippus, 294
ponceanus, 126
portlandia, 145
powellella, 194
Pseudocercyonis, 16
rapae, 144

rectilineata, 174
refusana, 284

ribearia, 266

Riodinidae, 37
sandraella, 205
sangoana, 172
Saturniidae, 42, 234, 238
Satyridae, 12, 41, 145, 150, 256
Schinia, 109, 177, 181, 274
separata, 177

Speyeria, 44, 150, 247
Sphingidae, 41
Strymonini, 87

tharos, 143

Tortricidae, 115, 284
toxeuma, 245

Urbanus, 294

Vanessa, 114, 147
Vanessinae, 140

verna, 286

Victoria, 169
virginiensis, 84
walsinghami, 181
watsonae, 169
wislizeniella, 209

Zetes, 142

Author Index

Bolte, 266

Bowden, 6

Brown, 152, 176, 240, 298
Brussard, 22

Byers, 124

Cannon, 150
Carcasson, 169
Clench, 80
Davis, 168, 247
Dominick, 84

302

Ebner, 73
Edwards, 84
Emmel, 12, 20
Ferris, 44

Fisher, 44

Ford, 155
Gatrelle, 143, 145
Gray, 65, 108
Greenfield, 278
Hardwick, 1, 109, 177, 181, 274, 287
Irwin, 83
Johnson, 216
Kaddow, 53
Kendall, C. A., 29
Kendall, Roy O., 29
Lafontaine, 299
Lehman, 150
Lighty, 239
Lindsay, 64
Manley, 146, 234
Martin, 215

Masters, 19, 84, 86, 149, 150, 256, 297

Mather, 147
Mattoon, 247
McFarland, 144
McGuffin, 262, 266
Miller, L. D., 12
Miller, W. E., 284

JOURNAL OF THE LEPIDOPTERISTS SOCIETY

Munroe, 142, 185
Nicolay, 87
Oliver, 143
Opler, 115, 194
Owen, 271
Palkuti, 150
Patterson, 222
Pechuman, 82
Peters, 114

Platt, 278

Pliske, 294
Rindge, 143
Rutkowski, 126, 137
Schwehr, 139
Sevastopulo, 80, 146, 293
Slansky, 294
Spencer, 247
Stary, 53
Steinhauser, 295
Steyskal, 264
Straatman, 58
Sullivan, 295
Tilden, 293
Toliver, 213, 246
Weave, All

Wood, 83
Wright, 68
Young, 223

NOTICE TO CONTRIBUTORS

Contributions to the Journal may deal with any aspect of the collection and study
of Lepidoptera. Shorter articles are favored, and authors will be requested to pay
for material in excess of 20 printed pages, at the rate of $17.50 per page. Address
all correspondence relating to the Journal to: Dr. D. F. Hardwick, K. W. Neatby
Bldg., Central Experimental Farm, Carling Ave., Ottawa, Canad.

Contributors should prepare manuscripts according to the following instructions;
failure to do so will result in unnecessary delay prior to publication.

Text: Manuscripts must be typewritten, entirely double-spaced, employing wide
margins, on one side only of white, 814 x 11 inch paper. Authors should keep a
carbon copy of the MS. Titles should be explicit and descriptive of the article’s
content, including the family name of the subject, but must be kept as short as
possible. The first mention of a plant or animal in the text should include the
full scientific name, with authors of zoological names. Underline only where italics
are intended in the text (never in headings). References to footnotes should be
_ numbered consecutively, and the footnotes should be typed on a separate sheet.

Literature cited: References in the text should be given as, Comstock (1927)
or (Comstock 1933, 1940a, 1940b) and all must be listed alphabetically under the
heading LireratureE Crrep, in the following format:

Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
1940a. Notes on the early stages of Xanthothrix ranunculi. Bull. So.
Calif. Acad. Sci. 39: 198-199.

Illustrations: All photographs and drawings should be mounted on stiff, white
backing, arranged in the desired format, for reduction to the page size (4% x 7
inches), including space for legends. The author's name, figure numbers as cited
in the text (where these are not intended to be engraved on the plate), and an
indication of the article’s title should be printed on the back of each mounted
plate. No charges are made to authors for line drawings or halftone (photographic)
figures, provided these are submitted in satisfactory condition for reproduction
without touch-up work. Figures, both line drawings and halftones, should be num-
bered consecutively in Arabic numerals. The term “plate” should not be employed.
Figure legends must be typewritten, double-spaced, on a separate page (not attached
to the illustrations), headed ExPLANATION OF FicuREs, with a separate paragraph de-
voted to each page of illustrations.

Proofs: The edited MS and galley proofs will be mailed to the author for
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Tables: Tables should be numbered consecutively in Arabic numerals. Headings
for tables should not be capitalized. Tabular material should be kept to a minimum
and must be typed on separate sheets, and placed following the main text, with the
approximate desired position indicated in the text. Vertical rules should be avoided.

Material not intended for permanent record, such as current events and notices,
should be sent to the editor of the News: Dr. C. V. Covell, Dept. of Biology, Univer-
sity of Louisville, Louisville, Kentucky 40208.

Memoirs of the Lepidopterists’ Society, No. 1 (Feb. 1964)
A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

by Cyrm F. pos Passos
Price, postpaid: Society members—$4.50, others—$6.00; uncut,
unbound signatures available for interleaving and private binding,

same prices; hard cover bound, add $1.50. Revised lists of the
Melitaeinae and Lycaenidae will be distributed to purchasers free.

ALLEN PRESS, INC. ap LAWRENCE, KANSAS
usr

CONTENTS

Brown, F. M. The “Arrowhead Blue,” Glaucopsyche piasus Boisduval
(Lycaenidae: Plebejinae) 2.0) ee 240-246

Hardwick, D. F. The life history of Schinia ligeae (Noctuidae) 274-277
Hardwick, D. F. The life history of Heliolonche pictipennis (Noctuidae) _. 287-292
Lighty, P. M. Neutron irradiation in Antheraea eucalypti Scott (Saturnii-

are) eal Na ee 239-240
Manley, T. R. Two mosaic gynandromorphs of Automeris io (Saturniidae) 234-238
Masters, John H. A note of Lethe anthedon borealis (Satyridae) —.___. 956-261
Mattoon, S. D., R. D. Davis and O. D. Spencer. Rearing techniques for

species of Speyeria (Nymphalidae) 247-256
McGuffin, W. C. Descriptions of larvae of two eastern species of Lygris

(Geometridae) | 22200 ee 262-264
McGuffin, W. C. and K. Bolte. Variation in larval colour patterns of Itame

ribearia (Geometridae) Eee 266-270
Miller, William E. Identity of Phaneta refusana (Walker) with description

of a new species (Tortricidae) __...._.__._..__..__.._ ee 284-287
Owen, D. F. Pupal color in Papilio demodocus (Papilionidae ) in relation

to the season the year 00 271-274

Platt, A. P. and J. C. Greenfield, Jr. Inter-specific hybridization between
Limenitis arthemis astyanax and L. archippus (Nymphalidae) —_. 278-284

Pliske, T. E. Notes on unusual species of Lepidoptera from southern Florida 294
Slansky, F. Jr. Danaus plexippus (Nymphalidae) attacking red-winged

black Dosa ipo 000 ac I al Sn 294
Sevastopulo, D. G. Remarks on “The use of net traps at Palawan, Philip-

Pines” °c yh ON pase NE er 293
Steinhauser, S. R. A simple method for preparing male hesperiid genitalia

for examination without dissection ... 295
Steyskal, G. C. On the grammar of the name Heliothis Ochsenheimer

(Noctuidae): 00 ho. SS 264-266
Sullivan, J. B. Captures of Erora laeta in North Carolina (Lycaenidae) __ 295-296
Tilden, J. W. Aguna claxon (Hesperiidae ) new to the United States ____ 293
Toliver, M. A record of Euphydryas anicia (Nymphalidae) in Oklahoma — 246
Young, A. M. Notes on gregarious roosting in tropical butterflies of the

genus Morpho 2.0). 00 ee 223-234

Book Reviews: Te) ce 297-299

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Volume 25

JOURNAL

of the

_ LEPIDOPTERISTS’ SOCIETY

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY

- Publié par LA SOCIETE DES LEPIDOPTERISTES
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

VOLUME 25

SUPPLEMENT l

A NEW GENUS OF HAIRSTREAK FROM
CENTRAL AND SOUTH AMERICA

(Lycaenidae, Theclinae)

BY S so, Nieolay

10 May 1971

THE LEPIDOPTERISTS’ SOCIETY
EDITORIAL COMMITTEE

D.F. Hardwick, Editor of the Journal
C.V. Covell, Editor of the News
S.A. Hessel, Manager of the Memotrs

Executive CouncIL (1971)

C.L. Remington (New Haven, Conn.), President

L. M. Martin (Prescott, Ariz.), President-elect

H.A. Freeman (Garland, Texas), Ist Vice-President

Julian Jumalon (Cebu City, Philippines), Vice-President

K.W. Philip (Fairbanks, Alaska), Vice-President

$.S. Nicolay (Virginia Beach, Va.), Treasurer

J.C. Downey (Cedar Falls, la.), Secretary

L. D. Miller (Sarasota, Fla.), Secretary-elect
Members at large (three year term): M. Ogata 1972

A.E. Brower 1971 E.C. Welling 1972

W.C. McGuffin 1971 Andre Blanchard 1973

Y. Nekrutenko 1971 R.B. Dominick 1973

B. Mather 1972 J.P. Donahue 1973

Be) Fe) Fert rel | ale

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formed in May, 1947 and formally constituted in December 1950,
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ee a ae

JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

Votume 25

SUPPLEMENT 1

A NEW GENUS OF HAIRSTREAK FROM
CENTRAL AND SOUTH AMERICA

(Lycaenidae, Theclinae)

Cole S& Se Nitcolays-USMCG (Ret...)
1500 Wakefield Drive, Virginia Beach, Virginia

ey:

Lia

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ae:
ws

INTRODUCTION

In a recent issue of the Journal of the Lepidopterists'
Society appeared a statement that the taxonomic status of
the South American ''Theclas'' is chaotic (Brown and Mielke,
1967). For those familiar with the tropical Theclinae, this
statement comes as no Surprise. Indeed, it may be expanded
to include most of the tropical hairstreaks of the New World.
The purpose of this work is to attempt to clarify the tax-
onomy of one small segment of the Theclinae (Strymonini) of
the American tropics.

At infrequent intervals, | have received small shipments
of hairstreaks from a few collectors in South and Central
America. Among these, three species were of particular in-
terest; each was represented by relatively good series and
included both sexes; each bore an unusual resemblance to the
other in size and wing shape, if not in color and pattern.
The three species were tentatively identified and placed to-
gether as a single group, although they were from two diff-
erent groups in Seitz (1920). A study of the male genitalia
confirmed the validity of the single group concept.

During the past six years | have collected four addi-
tional species of this new genus in the Republic of Panama,
the? Ganal Zone and in Colombia and Brazil. Recently, repre-
sentatives were found in the collections of the Smithsonian
Institution, the American Museum of Natural History, and the
Carnegie Museum. A single female of a new species was found
In a series of miscellaneous accessions in the British Museum
(Natural History). Analysis of these 10 species indicates
they belong to a distinct and narrowly defined genus, char-
acteristic in size, wing shape and general pattern. The
male and female genitalia are very distinctive. These, plus
other morphological characters, should make recognition of
any additional species relatively easy.

The original descriptions of many of the species treat-

ed herein are sketchy. Indeed, the descriptions may be
applied to any number of species outside this new genus.
Therefore, each is here redescribed and figured. The illus-

trations were made by the author; the photographs, with the
very able technical assistance of SSgt. Robert Prisler, U.S.
Marine Corps.

ACKNOWLEDGMENTS

My special thanks and appreciation go to Dr. John Downey,
University of Northern lowa, Cedar Falls, for his encoura-
ging critical analysis, many fine suggestions and excellent
technical guidance. To Gordon B. Small, Jr., Balboa, Canal

Zone, | am particularly indebted for his invaluable assist-
ance in all aspects of the field work, much of which made
this study possible. | am especially grateful to Mr. 1.G.

Howarth and Mr. G.E. Tite, British Museum (Natural History)
for their assistance and generosity in allowing me to exam-

4 Symbtopsts

ine and study the Druce and Hewitson type material. To Dr.
F.LH. Rindge, American Museum of Natural History, New York,
Dr. J.F.G.. Clarke, Dr. W.D,. Duckworth and Wi. D.. FiseiGiieeern—
sonian Institution, and Harry Clench, Carnegie Museum, my
thanks for their interest, helpful suggestions and complete
cooperation.

In Brazil, Dr. Keith S. Brown, Rio. de Janeigo sang ar.
Heinz Ebert, Rio Claro provided me with the opportunity to
study their large and interesting collections of Brazilian
Lepidoptera. To them go my thanks for their hospitality,
their many helpful suggestions and enthusiastic encourage-
ment. With Keith Brown | made an incredibly productive and
interesting collecting trip into the Mato Grosso. This
unique Opportunity to collect and work in such a remote area
is gratefully acknowledged.

SYMBIOPSIS Nicolay, new genus

Type species: Thecla strenua Hewitson, 1877

Hindwing with two tails, the shorter at end of Cuj, the
longer at end of Clu5. Postmedian line on underside Gitore—
wing ending abruptly at vein Cuy; without cell-end streaks
or bars on underside of fore- or hindwing. Postmedian Tine
of hindwing, bipartite, uneven, forming posteriorly a broad
angular "W" which terminates at 3A very close to inner mar-
gin; middle angle of "W" rounded, extending basad in inter-
space Cuo more noticeably than in other interspaces. Male
with no scent spot or other secondary sexual characteristics.
Frons with appressed scales and many intermixed erect bristle-—
like scales; eyes densly covered by short erect hairs; an-

tennae with a 15-segmented stalk, 12 segments on the club.

Male genitalia without a saccus; tegumen extended in a
horizontal, dorso-lateral plane, vinculum then curved sharp-
ly 90° toward ventral surface; falces viewed from ventral
surface, large, sharply angular, with terminal arms tapered
to a sharp point; viewed laterally, flattened dorsoventrally,
appressed rather close to uncus. Valvae long, 4/5 length of
vineulum, relatively narrow, completely separate, with in-
terior surfaces clearly grooved to retain aedeagus; aedeagus
Stout, evenly tapered to tip, posterior end curved dorsally
through approximately 60° in an even sweeping arc, with or

without a ventral keel, without cornuti or terminal teeth.

A new genus of Hairstreak 5

Female genitalia with ductus bursae a complex, stout
organ; dorsally composed of two lateral free-moving sclero-
tized elements, separated almost their entire length, fused
impo wa hollow, Sclerotized tube prior to entry into corpus
bursae ; ventrally composed of a semi-membranous pouch which
appears to hold the two dorso-lateral elements together.
Ostium guarded dorsally by the two rather heavily spined
@Geomsal plates of ductus bursae; ventrally formed by mem-
Branous juncture of seventh and eighth abdominal tergites.
Anterior @€nd of ductus bursae rather sharply recurved dor-
coilyaparoueh an arc of almost 180° just prior to entry into
mic corpus bursae; latter a simple, oblong, completely una-
Gerned sac, at least as long as or longer than the ductus
bursae.

The name Symbtopsts is an arbitrary combination of Latin
terms, but without special meaning. | consider it to be of
the feminine gender.

Symbtopsis Species bear a superficial similarity to many

new world tropical hairstreaks. The lack of scent spots and
other secondary sexual characteristics, the size, gro%nd
color and trace of the postmedian line on the underside of

the wings are features common to both Symbtopsts and the
genus Calystryma Field, 1967. Yet there are definite and
obvious generic differences between the two. Symbtopsts spe-
cies have no cell-end streaks on the underside of either
Wing; Calystryma species have the cell-end streaks on the

underside of both wings. The wing shape of Symbtopsts is
rounded and full; that of Calystryma, more angular, the apex
Of the primaries more acute, particularly in the male. The

male genitalia of Symbtopsis has no saccus, the falces are
large, sharply angular, the aedeagus is stout with a sweep-
ing dorsal curve, and without cornuti or terminal teeth; that
of Calystryma always has a saccus, the falces are rather
small, not sharply angular and the aedeagus is long, rather
slender, Usually straight and always with at least a single,
spine-like cornutus at or near the terminus. The female duc-
tus bursae of Symbiopsts is heavily sclerotized, divided
throughout most of its length into two lateral elements and
anteriorly recurved dorsally through an arc of nearly 180°,
prior to entry into the corpus bursae which is unadorned;
that of Calystryma is a simple sclerotized tube, not recurved
dorsally at the anterior end and with the corpus bursae orna-~
mented by two large, complex signa.

Both male and female genitalia are generically very
characteristic but do not offer a panacea for specific deter-
mination. The genitalia of both sexes show considerable vari-
dation within each species; e.g., the spines of the ostium
bursae vary both in size and number in females of the same

6 Symbtopsts

species. In this respect, most were found to be markedly
asymetric in the number and shape of the spines on each lat-
eral plate. The size and shape varies in much the same way

in the male genitalia of a given species, but to a lesser
degree. However, the combination of characters provided by
the genitalia of both sexes together with those of wing shape,
color and maculation make specific determination of both male
and female specimens relatively easy and accurate.

Most species of this genus are intensely local. Those
we have taken fly very rapidly along chosen paths or narrow
roadways skirting the sunny edge of usually heavily wooded
areas. Their rarity in collections jis understandable for
they are not readily attracted to flowers and rest only
briefly on leaves. In many cases, they must be taken on the

Figure 1. Symbtopsts hindwing; stylized sketch of underside
maculation pattern

A new genus of Hairstreak 7

wing. Their swift and erratic flight makes them difficult to
see, follow and net.

A stylized sketch of the undersurface pattern of the
hindwing of Symbiopsis is illustrated in Figure 1. Not all
species in the genus have all of the linear features shown,
nor do all species follow the exact pattern as illustrated.
However, the basie features of the genus are contained in
Figure 1 and the nomenclature for the specific macular fea-
tures is noted on the illustration and used throughout the
text and in the key.

Key to Symbtopsts species

] Underside of hindwing without red cubital spot........ 2
Underside of hindwing with red cubital S.BOib 2d brhvaven ce vats 3
2 Underside of hindwing without powder blue or grey-blue
spot in Cuy; underside ground color, greyish-white....
Serre ee OE od Sh, welt co ah lenttas (Druce)

Underside of hindwing with large grey-blue spots in

ieeemepaces Cu; and) Cu9......2% tanaits (Godman & Salvin)
3 Underside ground color light greyish-white; cubital
~rreeange red, Smabl...i:...65 .eese00 see ntppta (Dyar)

Underside ground color much darker, brown or brown-

ie ee eses on mee. St. Ge BBS e Ree be RS SOR D Ew ALES. 4
us Upperside of male and female forewing blackish-
ee ac ccla hs avr Mile ss he ee SNe e ER CORRS wee webu ee se 5

Upperside of male and female forewing with lustrous

EE ENG reac ch tag, oan shee s. 2,i6kk seu as ay 8. hig Gaeetind 0g 4 Sys, MER ainnel ines 6
5 Upperside of forewing and hindwing black-brown with

a faint blue-grey tint toward the wing bases; male

pees AAT Ke 28 2 RSs he a oS So Se puptlla (Draudt)

Upperside of male hindwing shining lustrous blue with

Wieeodark margins; female brown.........s. pentas Nicolay
6 Upperside wing color lustrous indigo-blue with wide,

Gavky margins...... Bc Ae re RHI wie le, at eck 1G A Ge i Wi Fz

Upperside wing color greenish steel-blue with narrow

ERICH I CEE FS Bocid ss & Geet w anim © wlel a) «(sap agfel SAS 20 wie Waele Se ie nee 9
7 Upperside of forewing mostly dark blackish-brown, the

blue confined narrowly in the disc along vein 2A......

REN Re ct As 2 Oe cod SR uw ka etna, ONG pennatus (Druce)

Upperside of forewing with the blue expanded and
Bate nmenenbo. the CEU a. Gi. cake sc tel eee ee eee ee 8

8 Symbtopsts strenua

8 Cubital spot of Cu, large, red, spilling into Space

M3, with a broad bar over the blue spot of Cug......
Sieltaa is lee et katte. otiakanabiey atanata “die va RaMoh a: he aan anaes lee smallt Nicolay

Cubital spot, of interspace Cu, shall, sauitmrerm
crescentics;> no red bar over the blue spot 228 i ae
Pe ten ota og bite eel a celle bes oe Baise ole aaa eI ise ara va

9 Cubital spot orange-red confined primarily to Cug,
the red bar: over the blue: spot; narrow... foie eee

Ss eae tee Nee Mala are an ok wean, ais etn Leen aia ene emenene panamensts (Draudt)

Cubital spot very large, deep red, spilling into
interspace Mater yen M3; a broad red bar over the
blue Spot Or’ Cura) 0. eee satatate ec ali ciate otal ora eee morpho Nicolay

SYMBIOPSIS STRENUA (Hewitson), new combination
Figure) 23 Plater! qFigie i andag)

Theela strenua Hewitson, 1877, Illustr. Diurnal Lepid; Ly-
caenidae, 1:207; 2, pl. 82, figs. 689, 69@ 0 Gmme ca:
Seitz, 1920 Gross-Schmett. der Erde, 5:788, pl. 156,
figs. i-9;)i-10.. Comstock and Huntington, 296gee0e0":
YY. Entiat SOC whe ip lereoar.

Type data: ''In the collection of W.C. Hewiltson,.agom
BireeaZau liar.

This species is very similar to many other tropical
hairstreaks. The original description is extremely brief and
overlooks many features that place strenua within this genus.
The species is redescribed as follows.

Male: Length of (forewings i2. 5 mm.

Upperside: Margins of forewings wide and ill-defined,
with indigo-blue color confined to discal third of wing,
adjacent to inner margin. Margin of hindwings about 1 mm
wide, expanded at apex, remainder of wing dull indigo-blue,
faintly iridescent; anal lobe spot small, black, centered
with orange-red; a marginal black line, inwardly edged in
white, beginning at the small tail, tracing OUteremaeermurO
anal lobe; fringes of same area tipped in white.

Underside: Ground color olive grey-brown; submarginal
line of forewing dark brown, lunular and broken at each vein;
postmedian line dark brown, distally edged in white, begin-
ning at costal margin and ending abruptly at vein Cuy. Post-
median line of hindwing dark brown, becoming darker towards
anal margin, distally edged in white, rather uneven, beginning
at costal margin, then forming a broad angular "W" before
terminating at inner margin; submarginal band composed of
dark brown crescents between each vein, vaguely bordered on
each side by light scaling; cubital spot between tails ydui

A new genus of Hairstreak )

med, smlail, crescentic and centered at distal edge with a
prominent black spot; the cubital red crescent as wide as
ground color between it and the white line proximal to term-
ie tene; the grey-blue spot adjacent in interspace Cuo
large, quadrate, with a small red bar between it and the
small black spot of anal lobe; no other red in interspace
~Cuy except as associated with anal lobe; a marginal black
line, inwardly edged in white, beginning at vein Mj] becom-
inespmuekesh at Cus, tracing outer margin to anak lobe.

Female: Length of forewing 12 mm.

Same as male except some variation in the amount of
Dilue on upperside of forewing; usually the pale blue color
Herkowmly cont ined to the area along inner margin, leaving
forewing almost completely dark brown; in early "spring
season” (October), Santa Catarina individuals, the blue of
forewing not confined to. inner margin, extending as a pale
wash far into. disc and cell, leaving a relatively narrow,
waepuely defined dark margin. The dark margin of hindwing
Seven byawider than in male. Underside as in male.

Symbtopsts strenua has in the past been confused with
Thecla ptstdula Druce, due probably to the Draudt (in Seitz,
Pee ewiistration of ‘'ptstdula; it is a good reproduction
of strenua. The series of two males and seven females from
which the present diagnosis was made was collected by Mr.
Fritz Plaumann in Nova Teutonia, Santa Catarina, Brazil, at
an altitude of 300-500 meters. The dates of capture are
predominantly January through March. Specimens from this
series were compared with the Hewitson type material in the
British Museum (Natural History).

A study of series of this species in the Brown and Ebert
collections provided material basis for a more accurate
appraisal of the geographical distribution. This species is
found in the forested mountain localities in the states of
Minas Gerais, Sao Paulo, Parana, Santa Catarina and to the
west in adjacent areas of Paraguay (Villa Rica). Most like-
ly, strenua is basically a southwestern species which reach-
es the northern limit of its range in Minas Gerais (Belo
Horizonte) via the tributaries of the Plate River system.

SYMBIOPSIS LENITAS (Druce), new combination
Figure 3; Plate | (Fig. 3 and 4)

Theela lenitas Druce, 1907, Proc. Zool. Soc. London, : 603,
Diy oo, tie. 5.0) Draudty tm: Selita, 1920, Gross=Sehmett.
der Erde, 5:788, pl. 156, fig. 1-8. Comstock and Hunt-
ineton, 1961, J. N. Y. Ent Soc., 697109.

Taspis vtolescens Spitz, 1931, Revista de Entomologia, Sao
Paulo, 1:48 (new synonymy).

10 Symbiopsts lenitas

Original description:

"Male. Upper side: fore wing uniform’ dull Drown;
paler towards centre of disc, inner margin very narrowly
dusted with blue. ‘(Hind wing pale violaceous Dive lace
costa, apex and outer margin rather broadly brown; abdom-
inal fold paler: brown. Cilia of fore wing Drowntenoaee
hind wing white. Underside ground colour uniform pale grey.
Forewing: a linear dark ultra-median band, outwardly bor-
dered with white, from the costa to the lower median ner-
vule, followed by a faint submarginal lunular line. A very
fine anteciliary dark line. Cilia yellowish grey, darker
at the tips... Hind wing: amedian linear band as ton tare
wing but more sinuous, commencing on the costa and reaching
to the abdominal margin, followed by a submarginal faint
lunular line which becomes more distinct towards the anal
angle. A black anteciliary line from the apex to the auaLl
angle. broadest at the base of the tails. Cilia patevarey.
Tails blackish tipped with white.

Female: Upperside uniform dull brown. Underside as
Gy Mes er

Expanse: 2 = 1/5 Aneh:

Habitat: Chapada Campo, Brazil (H.H. Smith); Paraguay
(Perrens).

Type, Mus. Godman.

A species with no brand or perceptible patch on the
fore wing and not allied to: any with which I am acquainted
but perhaps belonging to the group which contains 7. dindy-
Mus (Ce.

Mr. Smith's specimens were captured in January and
February."

The original description is excellent. This species is
one of the most easily recognized and distinctive of the
genus, made so by the lack of typical thecline markings on

the underside of the hindwings. There is neither cubital
red spot in interspace Cuy, nor blue spot adjacent thereto
in interspace Cuy. Instead, these are replaced by vague
patches of grey scales. The anal lobe spot is very smal}
and black.

Male: Length of forewing, 11'.5 ‘mm.

Female: Length of forewing, 11.5 mm.

These notes were made from a series of 12 males and 13
females collected by Mr. Fritz Plaumann in Nova Teutonia,
Santa Catarina, Brazil, at an altitude of 300-500 meters.
Most specimens were captured during the months of March
through June. A single female taken at Caviuna, Parana,

A new genus of Hairstreak 1]

Brazil in April is in the American Museum of Natural His-
tory. Specimens from this series have been compared with
the type series in the British Museum (Natural History), and
with the type of 7. violescens Spitz which is clearly a syn-
onym of lenitas.

[In the Brown and Ebert collections, | was able to ex-
amine an extensive series of this species, taken in numerous
localities in southern Brazil and Paraguay. Essentially,
lenttas seems to be a species of the wooded watercourses of
the Brazilian Planalto. Its currently known geographical
range includes the states of Minas Gerais, Goias, Mato
Grosso, Parana, Sao Paulo, Santa Catarina and adjacent lo-
calities in Paraguay (Villa Rica).

SYMBIOPSIS PUPILLA (Draudt), new combination
Eigure: As) Plate: ly (Fig. Sand 6)

Thecta pupttla Draudt, in: Seitz, 1920, Gross-Schmett. der
Prbeeemool, pl.. 158, figs. i-4, i=5. -Comstock and
nuseetarton., L963, 3. N. Y.s Ent. Soe. 71:46.

Original description:

",.. is above black-brown, towards, the base with a
Slight blue-grey tint, on the hindwing with a large red anal
Spot being bordered at first by black, then by white; be-
neath brown-grey, the lines like in the preceeding (Th.
panamensis) but finer; the submarginal lunae almost extinct,
from the upper median vein to the proximal margin distinct
as an entirely straight black line being removed far inward
and, Goeuching dents of the postdiscal band. The red spot of
the tail and the anal-spot extremely large, connected by a
bread red bridge, the black pupil of the spot of the tail
very small, strigiform."

There are no obvious differences between the male and
female of this species. Even the size differential (the
males average slightly larger) is more individual than sex-
ual. The phrase, ''... above black-brown, towards the base
with a slight blue-grey tint'' is an excellent capsule des-
cription of the upper surfaces of both wings. Based on
specimens examined during this study, puptlla can be further
characterized as follows.

Upperside: Anal lobe spot of hindwing small, rather
than large, centered with a sprinkling of orange-red; a mar-
ginal black line and an inwardly adjacent white line begin-
ning in interspace above the short tail at M3, tracing outer
margin to anal lobe.

Male: Length of forewing, 12 mm.

Female: Length of forewing, 11 mm.

2 Symbtopsts puptlla

Underside: Ground coior of both wings a light, silky
brown-grey, submarginal lunular line of forewings, faint
and. indistinct; postmedian line narrow, black, distally
edged in white, beginning just short of costal margin, end-
imgrau vei, uly, Postmedian line of hindwing brown-black,
distally edged in white, beginning at costal margin, uneven,
then forming a sharply angular "W" before terminating at
inner margin; center of "W" mark or semi-macule dark brown;
cubital spot between tails large, deep red with a small,
strigiform, black pupil at distal ‘center ) tired scalttaager
cubital spot often spilling into interspace’ Mey Diveseror
adjacent to: cubital spot, also: Warre> prosciitaniaie edged with
a bridge of red scales, reaching black spot of ana tore,
the. submarginal line of vague, indistinct lunae; marginal
line black, inwardly edged in white, tracing outer margin
from vein Mo to anal lobe, expanded slightly at €achivem
terminus; spot onranaiy Tobe) pilaek.

The expanded description and illustrations herein have
been taken from a series of five males and ten females col-
lected by Mr. and Mrs. Jorge Kesselring in Joao Pessoa,
Paraiba, Brazil. The dates cover the period December through
June; one female was collected on 31 October. In the Na-
tional Museum, Rio do Janeiro, Brazil are a male and female
of puptlla taken at Belem do Para and another male taken at

Amapa. Dr. Ebert's collection contains a series of puperila
taken in Paraiba. A female from the Oberthur collection in
the British Museum carrying the label ''Amazonas'' is placed

here. Major A. Bedford Russell, British Coldstream Guards,

collected a single female at Apoteri, British Guiana in
September 1963.

Available information (Bailey, 1947) indicates that

Draudt's types have been destroyed. It appears advisable
that a neotype be designated. A neotype female, BRAZIL,
Joao Pessoa, Paraiba, 6 June 1954, is therefore designated

from the series described above and placed in the National
Museum of Natural History (Smithsonian Institution), Wash-
ington, D. C. (USNM Type No. 71451).

Seitz' terse "Guiana to Colombia and Bolivia'' covers a
very large area. The currently known geographical distri-
bution for puptlla is rather restricted, being confined to
the Guianas and the adjacent northeastern coastal area of
Brazil as far south as the state of Paraiba.

SYMBIOPSIS SMALLI Nicolay, new species

Figure ' 53) Plater Tl) (Crige.) 42 63eccmen a)
Male: Length of forewing, 13° 4 1 mm; ‘holotype. serum
Upperside: Forewing margin broad, black-brown, reach-

ing well into discal cell; the shining dark blue confined
to lower one-third of wing along inner margin, extending

A new genus of Hairstreak 13

past vein Cuy into cell. Hindwing margin narrow, black-
brown, slightly expanded at apex; remainder of wing shining
dark blue; anal lobe spot black, narrowly edged above and
below with pure white; a narrow, vague,white marginal line
feom smell tail at Cuz. to anal lobe.

Underside: Forewing ground color dark brownish grey;
Submarginal line vague, indistinct; postmedian line almost
straight, black-brown, distally bordered with a few white
scales, extending from near costal margin, terminating at
Cu2. Hindwing ground color dark brownish grey; postmedian
line uneven, black, distally edged in white, cleanly broken
inward at vein M3, terminating at inner margin after trac-
ing a broad deep "W"; submarginal line composed of vague,
light grey crescent-shaped lunae; a black marginal line,
edged basaly in white, beginning at vein M], becoming broad-
er and more definitive at cubital red spot between tails;
eubital spot orange-red, large, with a large strigiform
Diack pupil at distal center, spilling unevenly into adja-
cent interspace M3; a broad red band proximally borders the
large grey-blue spot, the two colors almost of the same
Weeaems anal lobe spot black, with proximal and,distal mar-
gins white, and with a connecting red line running costad
EG ered bar of Cu5; a-rudimentary black spot at basal margin
Suetoreesblue spot in) Cuo, touching the red bar’.

Female: Length of forewing, 12 mm.

Upperside: Markings and ground color similar to male;
the dark shining blue area less brilliant in forewing, re-
stricted to a small discal and basal area adjacent to inner
margin along vein 2A. Hindwing margins narrower, the dark
blue thus more extensive in the discal and basal area.

Underside: Ground color brownish grey with a slight
ieseee- ail linear markings Similar, to male, heavier,* the
red cubital spot expanded into interspace M2.

Holotype male: PANAMA, Los Rios, Canal Zone, 10 Decem-
ber 1967. Allotype female: same locality, 17 December 1964.

Paratypes, 41 male (m) and 7 female (f), as follows. From
Eeneoaneee smalls collection: Los Rios; €.Z., | m, 31 Oct.,
Seminowv. 66m e.b3 Nove, 1, myol4 Nov. 196531 nm, bec. ,
Mmmeaneeiec. lam, ..0,Decw, 1 m,.l2 dec, I amge13 Deetyylom,
nompcica = G64. 3m, 20 Dec. 1967s. Kem, 12 Febusrl omy 23 )Maneh
1968; Ft. Sherman, C.Z., 1 m, 21 May 1966; Cerro Campana,
Panama.Province, 1 m, 10 Jan. 1968; 1 f, 29 May 1964; Turri-
wWoae Cartage Prov., Costa Rica, ) f, 14Julye1965...From
Eernicolay collection: .Los Rios, C. Z2., 1m, bh Dee. 13 om;
Pebec anes) the, Ib Dec, 2 my S Décinl 967 7 lem Dees! om;
Pomece: slom, 24 Decw, 1964; 2 m, 3-f, 6 Jan. 1969; Colon
SameacRita).Rep~.of Panama, | m;5 Jane, 1 f, 4 Jans, 07 Ff,
LOjJdan. 1969:

The holotypeand allotype will be deposited in the Nation-

14 Symbtopsts pentas

al Museum of Natural History (Smithsonian Institution),
Washington, D. C. (USNM Type No. 71452). Paratypes will be
deposited in the American Museum of Natural History, New
York, the Carnegie Museum, Pittsburgh, and the British Mu-
seum (Natural History), London.

Symbtopsts smatlt closely resembles S. tanave oni the
upper surface. The basic blue and dark marginal colors are
alike in both; the size and shape of the adults of both
sexes are similar. However, the maculation of the under
surface of the wings is strikingly different. Here, smallt
closely resembles puptlla in the clarity and color of the
linear markings.

Thus far, smallt has been found in the Republic of
Panama, Costa Rica and Colombia, with a known altitude rang-
ing from sea level to 2000 feet. Its primary flight period
appears to be the winter months of November through February
although it has been taken in almost all seasons. Examined,
but not included in the type series, was a male collected in
Jaque, Darien Province, Panama in April and a male in the
British Museum, taken in Cundinimarca, Colombia.

It is a distinct pleasure to name this beautiful insect
after my good friend and collecting associate, Gordon B.
Small, Jr. who collected most of the specimens.

SYMBIOPSIS PENTAS Nicolay, new species
Figure 6:° Plate {Itt (Fig: 915 °2,453° ands)
Male: Length of forewing, 12 mm 2 1 mm; holotype wee ol.

Upperside: Forewing dark brown with.no blue scaling.
Hindwing opalescent violet-blue in the disc with wide (2-3
mn) dark brown margins, widest at apex; an inconspicuous
white submarginal line, beginning just above the small tail
at M3 tracing wing margin to anal lobe; latter small, black
with intermixed bronze colored scales, proximally and dis-
tally white fringed.

Underside: Forewing ground color light brown-grey;
Submarginal lunular line faint, dark brown, broken at each
vein; postmedian line dark brown, faintly and distally out-
lined with light scaling, ending’abruptly at vein Cus. itee—
wing ground color. the same brown-grey; submarginal line faint,
dark brown crescents, separated at each vein, distally and
proximally edged in faint light scaling; postmedian line
dark brown, distally edged in white, beginning at costal
margin, rather uneven, becoming black, forming a broad angu-—
lar "W" beginning at Cu, and ending at inner margin; cubital
Spot at Cu, small, orange-red, the black pupil distally cen-
vered ,; prominent; biue) spotwoL Cup proximally bordered by a
narrow bridge of red scaling beginning at black anal lobe
spot, not reaching vein Cus; semi-macule dark brown; a mar-

A new genus of Hairstreak BS

ginal black line, expanded at each vein and inwardly edged
in white, beginning at vein My and ending at anal lobe.

Female: Length of forewing, 11 mm + 1 mm; allotype,
i 1° mm .

Upperside: Forewing and hindwing dark brown with no
trace of blue; anal lobe spot black; marginal white line
from vein Mz very faint.

Underside: Same as in male.

Baborype mate: BOLIVIA, Cuatro Ojos, Nov. 1913. Allo-
type female: Las Juntas, Bolivia, Dec. 1913. Paratypes,
3 male, 5 female, as follows. In the Carnegie Museum: 1 m,
Paeatuneas, polivia, Nov. 1913; 1] m, Cuatro Ojos, Bolivia
famedabe)- 3. f, Las Juntas, Bolivia, Dec..1913, 1 £, Cuatro
Ores. Bolivia, Nov. 1913. In the Smithsonian Institution:
Pao agdiny +t, Peru, no date. W.D. Field in his discussion
of Calystryma malta Schaus, refers to a fourth specimen of
the original Schaus series, a female, as not belonging to
Calystryma. I|t is this female that is here designated a
paratype.

The holotype and allotype are deposited in the Carnegie
Museum, Pittsburgh, Pennsylvania.

The complete lack of blue on the upperside of the fore-
wings in the male makes this species relatively easy to dis-
tinguish from other members of the genus. However, the
female is almost indistinguishable from the female of penna-
tus except by examination of the genitalia. Both are dark
brown on the upper wing surfaces and very similar beneath.
The most consistent difference between this species and
Ngereecus, 1s in; the black pupil of the cubital spot;3,.in pen-
Po2ofhis Spot 1S round and prominent, in pennatus) it. ts) stri-
giform, ill-defined and almost obsolete in some specimens.
In the genitalia, particularly that of the male, there are
consistent and easily recognized differences as noted -in the
appropriate figures.

| have seen no specimens of pentas other than those in

the type series. The known range of this species is Bolivia
and Peru.

SYMBIOPSIS PANAMENSIS (Draudt), new combination

Eirgure 2 2 hate ht kh baa1> cand, G)

Thecla panamensts Draudt, tn: Seitz, 1920, Gross-Schmett. der
Erde, 5:801, pl. 158, figs i-l, i-2. Comstock and Hun-
Penerc Ons, 1 62,4 disp diem Yoo HMtr, COC ce. 7 Ol: LOTS

Original description:

16 Symbtopsts panamensts

t.. Ls. the most similar’ to ptetdula; but 10 feeaneue
of a light greenish steél—blue, with a strong Iustres eee.
margins of the. same extent as in pisidula; the fematewiewo:
a duller grey-blue. Beneath more purely grey, with a slight
lustre, the linear marking stronger, particularly the white
bordering more pronounced, the submarginal lunae more dis-
tinet, the red spot of ‘the tail very ‘large, so> 2S” (iemieseae
blue spot beside it.”

The sexes are similar in appearance, and the descrip-
tion may be elaborated as follows.

Upperside: ~ Greenish steel-blue Justre accentuaveamay
the narrow black margins, more so than in any other species
except morpho; margin rather sharply defined in forewing of
male, rather vaguely defined and slightly broaderiane tease.
black. marginal color almost filling primary wing Gell aan
female. Dark margin of hindwing consisting of thin line,
slightly expanded at apex, the same greenish steel-blue
filling entire hindwing in both sexes; anal lobe spotypdack
set within a sprinkling of white scales.

Underside: Submarginal line of forewing of the same
lustrous grey as basic ground color of both wings but Garker
and distinct, running from subapical area to tornus; post—
median line very dark grey, narrow, distally edged in white,
following the same curve as the submarginal but ends abrupt-
ly at vein Cuo’”. ‘Postmeditan of Nindwing a cConsprlceveus is
black line, distally edged in white, sliphtly broken? @e270—
ning at costal margin, then forming a broad "W" prices
termination at inner margin; semi-macule black; submarginal
lunae proximally shaded in light grey; marginal line from
M, to anal. lobe;black, inwardly edged in pure white; cubital
Spot between tails orange-red, large, with a Strigiform
black pupil at distal center; a very narrow red bar proxim-—
ally bridging. a large light blue spot adjacent in @snvereaoee
Cua, almost reaching the small black spot of anal lobe;
fringes white-tipped between the short tail at Cuj and anal
lobe.

The underside pattern of panamenstis closely resembles

strenua. All markings are similar but in panamensts are
more definitive, the spots larger and the ground color grey-
ish with a slight lustre. Males and females are alike, with
the ground color of the females generally lighter, almost

brownish rather than grey.

The above observations were made from specimens which

Gordon small and | collected at Potrerillos, Chi rrqauieerer-
ince, in-western Panama; dates of capture were in the months
of December, January, February and July. In addition, Gor-

don Small has taken a single specimen at Villa Neily in the
adjacent province of Puntarenas, Costa Rica in July.

Available information (Bailey, 1947) indicates that
Draudt's types have been destroyed. It appears adviseable

A new genus of Hairstreak Wy

that a neotype be designated. A neotype, male, PANAMA, Po-
trerillos, Chiriqui Province, 11 Feb. 1966, is therefore
designated from the series described above and placed in the
Nattonal Museum of Natural History (Smithsonian Institution),
Washington, D. CC. (USNM Type No. 71453).

SYMBIOPSIS MORPHO Nicolay, new species
Figure ©, Plate 111 (Fige 5S ands&Y
Male: Unknown.
Hemale: Length of forewing, 15 mm.

Upperside: Forewing costal and outer margins black,
narrow with a wide expanse of brilliant greenish steel-blue
Corocertiiane cell, disc, basal and most of inner marginal
area. Hindwing the same shining morpho-like blue; outer
margin a thin, black line expanded at apex; abdominal fold
Gdamemetey: anal lope spot large, biack, narrowly fringed in
white.

imG@ercisnde- Horewane eround color silky Might brown;
supmareinal line darker brown, faint but distinet; postme—
Gian line narrow, blackish-brown, distally edged in white,
ends at vein Cu2. Hindwing ground color light brown; post-
medtanetane Sharply black, distally edged in pure white, dis—
placed outward at interspace. Sc + R1, broken inward at vein
eee eeritnaLing at inner margin after tracing a broad "W";
Submarginal line dark brown, consisting of crescent-shaped
imap suadty and proximally edged in light scaling; cubi-
Vm pOrmeii Cun very large, deep red, proximally spilding in-
Demaeniiacemtu space, reaching vein Mas the black pupil centered
gueisval cdpe dargce, strigiform; blwe spot of Cua capped by
a rudimentary black spot, completely bridged proximally by a
“wea bar Extending from cubital spot to the large black spot of
enciehobie Lavver proximally and distally framed in white.

-Holotype femate: EQUADOR, Paramba, 3500 ft., March
1897, dry season (Rosenberg); in the British Museum (Natural
History), London.

This is the largest and most lavishly colored Symbtop-
sts. The upper surface is of the same shining steel-blue
lustre of panamensts, the dark margins similarly narrow.
Beneath, morpho is a rich brown color, rather than greyish.
[t is more heavily marked than any other species of this
genus, | suspect that the male, when found, will be larger
than the female, with narrower dark margins on both winas
and a larger expanse of brilliant blue color on the upper
surfaces.

In response to my request for specific information on
tne locality Paramba, Mr. Tite of the British Museum very
kindly provided the following quote from a paper by Dr.
Hartert dealing with the birds collected by W.F.H. Rosenberg:

18 Symbtopsts pennatus

'Paramba is a farm on the western bank of the river Mira.
Its elevation is 3500 feet, and itis still in’ the» forese
region, but the open country commences two or three miles
higher up the Mira...The city of Ibarra two days ride from
Paramba, and about the same distance from Quito..."

SYMBIOPSIS PENNATUS (Druce) new combination
Figure 9; Plate IV (Fig. 1 and 2)

Theela pennatus Druce, 1907, Proc. Zool. Soc. London,
7617-18. Draudt in: Seitz, 1920, Gross-Schmett. der
Erde, 5:800. Comstock and Huntington, 1962, J. N. Y.
Ent. Soe.) fOrtoer

Original description:

“Male. Allied to Tf. anthora Hew. Upper side uniform
dark brown, with inner margin of fore wing narrowly, and
discal and basal areas of hind wing shining opalescent indi-
go-bDlue.. Under side; ground colour duller; the linear bands
narrower, placed further in from the margin and in the hind
wing not inwardly bordered with red. The upper red marginal
spot with the black pupil of Tf. anthora is absent, and the
large red spot contained in the angle of the ultramedian
band is replaced by a black, smaller spot.

Female. Upper side uniform dull brown; underside as in
male.

Expanse, 1 .1/5,,.1 1/10 ineh..
Hab. Amazonas: Tapajos, Ega, Para (Bates).
Types, Mus. Godman."

The brevity and accuracy of Druce's original descrip-
tion needs little elaboration. The anal lobe spot on the
upper hind wing is black, centered with sparse orange-red;
the marginal black line from vein M3 to the anal lobe is
inwardly edged in white.

On the underside the cubital spot of the hindwing is
relatively small, orange-red and pupiled by a vague, ill-
formed almost obsolete black spot. The red bar, beginning
at the black anal lobe spot is narrow and, although arched
over the proximal edge of the blue spot of Cuyg, is incom-
plete and does not reach vein Cu9.

The male genitalia of pennatus has a ventral keel on

the aedeagus. lt is not obvious and is confined to the pos-
terior end. The genitalia of tanats and ntppta are similarly
equipped.

Male: Length of forewing, 12-13 mm.

A new genus of Hairstreak 19

Female: Length of forewing, 11-12 mm.

As stated by Druce, pennatus is certainly a distinct
species, but it is not related closely to 7. anthora Hew.
tieedectern Of lines and color it ts ‘most similar to pentae,
the females being almost identical. However, the genitalia
Feved! a close relationship to tanate and nippta, both of
which possess macular patterns beneath which are distinctly
different from pennatus. The most consistent difference
between pennatus and other species of similar appearance is
Mee Didcek pupil of the cubital: spot. In! pennatus, itis
small, ill-formed, almost obsolete, while in pentas and
others, it is prominent. The narrow blue scaling along
vein 2A in the forewing will readily separate the males from
pencade, its closest mimic.

The geographical range of this species is large, taking
in much of the vast expanse of the Amazon basin. The pres-
ent data adds Colombia, Peru and Bolivia to the political
regions given by Druce. Recorded altitudes are from near
sea level to 450 meters.

In addition to examination of the type series in the
British Museum, | have studied a male collected in Ega (Bra-
Ziv a series of four males and three females. from the Rio
Surutu and Portachuelo in eastern Bolivia, and a male from
Manacapuru, Amazon River, in the Carnegie Museum, and a
male and two females which |! collected on the Rio Bodoquero,
Caqueta, Colombia, in January.

SYMBIOPSIS TANAIS (Godman & Salvin), new combination
Figure 0; Plate IV (Fig. 3 and 4)

Theela tanais Godman and Salvin, 1887, Biologia Centrali-
imectieana., Lepid.—-Rhop. , 2:623)3:pl.,.. 55, figs 27,20.
Draudt tn: Seitz, 1920, Gross-Schmett. der Erde, 5:800,
peso, fie, 2-5. “Comstock and Huntington, 1964
Woes nes OCs sg Te riiede,

This is one of the most distinctive and easily recog-
nized species in the genus. The original description was
given in Latin, and the species may be characterized as
follows.

Male: Length of forewing, 13-14 mm.
Female: Length of forewing, 11-13 mm.

Upperside: Similar to smalit, differing as follows:
center of the anal lobe spot in tanazs orange-red, in
smallt black; blue scaling of forewing more restricted to
inner margin (never reaching vein Cu2) in tanatis, extending
Daeo vein Cud into the cell in emaliz.

20 Symbtopsts tanats

Underside: Ground color light grey-brown. The submar-
ginal and postmedian lines of forewing both faintly marked;
dark brown scaling filling semi-macule and distal to post-
median line in Cu, 3 the large powder-blue spot between tails
in interspace Cu, and an adjacent spot of the same (to
sligntly larger) size and color in interspace Cup provide the
most distinctive identification feature of this species,
anal lobe spot black, with a narrow red bar (the only red
color on the underside) between it and the blue spot adja-
cent; an obvious black marginal line, inwardly edged in
white and expanded at each vein terminus, beginning at vein
Mo, ending at anal lobe.

Male and female genitalia following basic pattern for
genus except aedeagus with a ventral keel.

Many of the specimens from which these observations
were made were collected in the Republic of Panama and the
Canal Zone by Gordon Small and I. Altitudes from which the
Panama specimens were taken range from sea level to 2500
feet. Specimens in Gordon Small's collection were taken as
follows: Cerro Campana, Panam Prov., December; Cocolij7e e.,
November; Madden Dam, €.Z., Aprils Summit, €.Z., Aprimteed
rather worn female, from Victoria, Caldas, Colombia, in
August. In the Nicolay collection: Cerro Campana, December,
January and February; Summit in April; Madden Forest in May
aNiGoPa nia, C Zc ced ieeniane.

There are four males in the collection of the Smithson-

ian Institution, all from various localities in Costa Rica.
One altitude notation reads 2500-3500 feet. The dates are
Nov. '06, Feb. and March. In the American Museum of Natural

History are three males and six females all collected by
H.H. and F.M. Brown at Turrialba, Costa, Rica on 24 amduem
June. All specimens noted above agree with the types loca-
ted in the British Museum (Natural History), London.

Godman and Salvin restricted the range of tanats to the
state of Panama; Seitz added the Rio Dagua in Colombia.
Symbtopsts tanats has thus far been taken in Costa Rica,
Panama and Colombia.

SYMBIOPSIS NIPPIA (Dyar), new combination
Figure lle Plate IVe(Fige 5 and 6)

Theela nitppta Dyar, 1918, Proc. U. S. Natl. Mus .5°55eeeee
Draudt in: Seitz, 1920, Gross-Schmett. der Erde, 5:824.
Hoffmann,1940, An. Inst. Biol., 11:774. Constoek uae
Huntington, 1964. /5)' NewYio ene Ssoes.h6Scavo.

Original description:

"Fore wing blackish, shaded with light blue on the ba-
Sal third below cell and in cell to its end. Hind wing blue

A new genus of Hairstreak 21

hoevern oO: fringe white:*a black terminal line; tail at

vein 2 long, white margined and tipped; tail on vein 3 short,
white; fore wing with faint whiter outer line, dislocated
aeeene veins. Hind wing with the outer line slender, black-
ish, edged without by white, forming a shallow W from vein
BeeeO Marein;s a faint submarginal lines; a red spot in the
interspace 2-3 with outer black center; a black and red

speck at tornus. Expanse, 25 mm.

Type. —— Female, Cat. No. 21201, U.S.N.M.: Siterra de Guerr-
ero, Mexico, January, 1911 (R. Muller)."

The male, which Dyar did not describe, is similar to
the female on the upper surface and identical on the under-
surface of the wings. On the upperside the blue of the
forewing is not as extensive as in the female, being con-
fined to the discal and basal area of the inner margin; the
hindwing has a wider dark margin (3mm at the apex) and the
blue of the discal and basal area is darker and more intense.
The anal lobe spot is red and a marginal black line with an
Hnnerewhite Line begins at vein M3 and ends at the anal lobe
spot.

lt is worthy of note that the two species with the
lightest ground color on the underside of the wings occur at
the southern and northern extremes of the known geographical
distribution of the genus, lenttas from southern Brazil and
Paraguay and ntppta from the state of Jalisco, Mexico. Both
are easily recognized by the greyish-white ground color upon
which the linear markings are rather faintly contrasted and
by the *'W'' mark of the postmedian line shallow rather than
angular. Dyar's "black and red speck at the tornus" refers
th the orange-red bar that lies between the small black
Saetrar the anal .lobe and the blue spot of Cu,. tt is areh-
ed partially over the proximal margin of the febeer. The
cubital spot is small, orange-red with a yellowish proximal
margin and centered at the distal edge with a prominent black

pupil.

As in tanats and pennatus, the male aedeagus of nippia
has a ventral keel. Otner small differences in the genitalia
can be noted in the appropriate figures.

I

In the American Museum of Natural History are four males
and five females from the C.C. Hoffmann collection taken in
April at Colima, Colima, Mexico. In the Carnegie Museum is
a male taken’ by R.G. Wind in May at Comala, Colima, Mexico.
The recorded range of ntppta is the west coastal area of
Mexico from the state of Jalisco south to Guerrero.

22

LITERATURE CITED

Bailey, J.W., 1947. Report on the status of the entomologi-
cal collections in certain European museums. Ann. Ent.
Sods Amen, 405203 24/2

Brown, K.S. and 0.H.H. Mielka, 1967. Lepidoptera of the
Central Brazil Plateau. I!. Preliminary list of Rhop-
alocera (continued): Lycaenidae, Pieridae, Papilioni-
dae, Hesperiidae. J. Lepid. Soc., 21:151-168.

Comstock, W.P. and E.I. Huntington, 1958-1964. An annota-
ted list of the Lycaenidae (Lepidoptera: Rhopalocera)
of the Western Hemisphere. J. New York Ent. Soc.,
vols. 66-72.

Draudt, M., 1916-1924, Lycaenidae. tn: Seitz, Die Gross-
Schmetterlinge de Erde. Stuttgart., 5:739-831, pls.
144-159, 193 (pps 788,801, pls: 156, 158;01saam

Druce, H.H., 1907. On Neotropical Lycaenidae, with des=
criptions of new species. Proc. Zool Soc. London,

1907:566-632 (:603, 617-618).

Dyar, H.G., 1918. Descriptions of New Lepidoptera from
Mexico. Proc. U. S. Natl. Mus., 54:335-372 (Lycaeni-
dae: 336-338).

Field, W.D., 1967. Butterflies of the new genus Calystryma.
Proc. Us. S.)NatlooMusey 123. (ne. 361.1): =a ee Gee

Godman, F.C. and 0. Salvin, 1887-1901. Biologia Centrali-
Americana. Insecta. Lepidoptera-Rhopalocera. London,

vol. 2: 1s782 bp. #62, Wea, <

Hewitson, W.C., 1863-1878. Illustrations of Diurnal Lepid-
optera. Lycaenidae... London, 121-228 (p. 0207) nnegeue
2. pls.) 1-92 (pls 8229 figs. 689 “and 690", (agai

Hoffmann, C.C., 1940. Catalogo systematico y zoogeografico
de los Lepidopteros Mexicanos. Primera Parte, Papili-
onoidea. An. Inst., Biol «,.11 9639-739 (p17 eae

Symbtopsts strenua 23

oH
ae

sere

SZ Sian

Figure 2. Symbtopsts strenua (Hewitson). a, ventral
view of female genitalia with 4vipositor and tergites remov-
ed; b, lateral view of female genitalia without ovipositor,
with 8th tergite; c, ventral view of male genitalia with
valvae and aedeagus removed; d, ventral view of valvae; e,
lateral view of male genitalia with aedeagus removed; i Tat=
eral view of aedeagus.

>
°
I

2h

Symbtopsts lenitas

Figure 3. Symbtopsts lenitas (Druce). a, ventral view
of female genitalia with ovipositor and tergites removed; b,

lateral
and 8th
vae and
al view
view of

view of female genitalia without ovipositor, with 7th
tergites; c, ventral view of male genitalia witha.
aedeagus removed; d, ventral view of valvae; e, later-
of male genitalia with aedeagus removed; f, lateral
aedeagus. ic

Symbtopsts puptilla 25

Figure 4. Symbtopsis pupilla (Draudt). a, ventral view
of female genitalia with ovipositor and tergites removed;
b, lateral view of female genitalia without ovipositor, with
8th tergite; c, ventral view of male genitalia with valvae
and aedeagus removed; d, ventral view of valvae; e, lateral
view of male genitalia with aedeagus removed; i, Tateral View
of aedeagus.

26 Symbtopsts smallt

Figure 5. .Symbtopsts smalli Nicolay. a, ventral view
of female genitalia with ovipositor and tergites removed;
b, lateral view of female genitalia without ovipositor, with
7th and 8th tergites; c, ventral view of male genitalia with
valvae and aedeagus removed; d, ventral view of valvae; e,

lateral view of male genitalia with aedeagus removed; f, hat-
eral view of aedeagus.

Symbtopsts pentas 24

Figure 6. Symbtiopsts pentas Nicolay. a, ventral view
of female genitalia; b, lateral view of female genitalia
without ovipositor, with 8th tergite; c, ventral view of male
genitalia with valvae and aedeagus removed; d, ventral view
of valvae; e, lateral view of male genitalia with aedeagus
removed; f, lateral view of aedeagus.

28 Symbtopsts panamensts

Figure 7. Symbtopsts panamensts (Draudt). a, ventral
view of female genitalia with ovipositor and tergites removed;
b, lateral view of female genitalia without ovipositor, with
bth tergite; c, ventral view of male genitalia with valvae
and aedeagus removed; d, ventral view of valvae; é Va tetral
view of male genitalia with aedeagus removed; f, Tatermat
view of aedeagus.

Symbtopsts morpho

29

Figure 8. Symbtopsts morpho Nicolay. a, lateral view
of female genitalia without ovipositor, with 7th and 8th
tergites; b, ventral view of female genitalia without ovi-
posi or, with part of 8th tergite.

30 Symbtopsts pennatus

Figure *9.) Syumbtopersipenmar ver (DrMeell. are
of female genitalia with ovipositor removied, w
8th tergite; b, lateral view of female genitali

ovipositor, with 8th tergite; c, ventral view of make gent=
talia with valvae and aedeagus removed; d, ventral view of

valvae; e, lateral view of male genitalia with aedeagus in
BOeLU.

>
ith part of
a without

ventral view

Symbtopsts tanats 31

Figure 10. Symbtopsts tanaits (G & S$). a, ventral view
of female genitalia with ovipositor and tergites removed;
b, lateral view of female genitalia without ovipositor, with
8th Leng tes ¢j.. variation, in spines of esStium;, d, wenthal
view of male genitalia with valvae and aedeagus removed; e,
Ventral view of valvae; f, lateral view.of male genitalia
with aedeagus removed; g, lateral view of aedeagus.

32 Symbtopsts ntppta

of female genitalia with ovipositor and tergites removed;
b, lateral view of female genitalia without ovipositor, with
Bth tergite; c, ventral view of male genitalia with valvae
and aedeagus removed; d, ventral view of valvae; e, lateral
view of male genitalia with aedeagus removed; *, Vatepaa vier

of aedeagus.

Figure ll. Symbtopsts nippta (Dyar). a, ventralewienm
i

Peat eo

33

Figures 1, 2, Symbtosts strenua
Teutonia, Santa Catarina, Brazil; 3
male, Nova Teuton ae

(Draudt) (neotype) male, Joao Pessoa,

(Hewitson) male
el Posh
Santa Catarnia, Brazil

.
>

Paraiba

» Nova

Lenttas (Druce)

5, 6, S. pupilla
> Shake

=

a

|
‘ # J
% ‘e RY Ua)
i Je Maes
i a '
t ‘
\ iy ahs
ia is
¥ is * Ny,
* Ne
7)
’ ; ¢ r
iat eh Sete AN ea RORY
~*~ 74 nd { ;
. ave Ae Nr a
meth p ‘
4 , * ‘
4 rat nt
acs
at ey heed
~ ‘ 4 (een
, Na"s
‘ 4 ‘
Pt
Eye,

35

PLaTe I]

Figures 1, 2, Symbtopsts smalli Nicolay (holotype) male,
Los Rios, Canal Zone, Panama; 3, 4, S. smalli (allotype)
female, Los Rios, Canal Zone, Panama; 5, 6, S. panamensis
(Draudt) (neotype) male, Potrerillos, Chiriqui Prov., Panama.

a7

Pioame l

Figures 1, 2, Symbtopsts pentas Nicolay (holotype) male,
Cuatro Ojos,. Bolivia; 3, 4, S. pentas (allotype) female, Las
Juntas, Bolivia; 5, 6, S. morpho Nicolay (holotype) female,

Paramba, Equador.

’

AOL aaa NS

Plate IV a

ymbiopsis pennatus (Druce) male, Ega,
ae hes 6. tanate MG. & So diomahe, Cerne
6, S. nippia (Dyar) male, Comala, Colima

Figures I, 25 6
"Amazonas", Brazi
Campana, Panama;
Mexico.

ws
ww

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VOLUME 25
SUPPLEMENT 2

Met OSYSTEMATIC STUDY OF TWO SPECIES OF
CALLOPHRYS (CALLOPHRYS) IN CALIFORNIA

(Lycaenidae)

by Glenn Alan Gorelick

10 May 1971

‘

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JOURNAL OF THE LEPIDOPTERISTS’ SOCIEiY

VoLume 25

SUPPLEMENT 2

A BIOSYSTEMATIC STUDY OF TWO SPECIES
OF CALLOPHRYS (CALLOPHRYS)
IN CALIFORNIA
(LYCAENI DAE)

Glenn Alan Gorelick

Citrus College, Azusa, California

TABLE OF CONTENTS

Introduction
Acknowledgements
Historical Review
Methods and Materials
Geographic and Ecological Distribution
Comparative Morphology
Evaluation of Characters: Callophrys dumetorum
and Callophrys viridis
Evaluation of Characters: Callophrys dumetorum
perplexa
General Biology
Biology of Callophruemaumetoenum ii. “425g ae
Adult Behtavi ons « cae.
Life History
Biology of Catlophrys virtdts
Adult Behavior
Life History
Results and Conclusions
Literature Cited
Nepenai (Materials Studied)

Figures and photographs

INTRODUCTION

The genus Callophrys Billberg is represented in North
America by six subgenera separable on the basis of male gen-
italia (Clench, 1961). These subgenera are Xamia (one spe-
cies), Sandia (one species), Inetsalia (eight species), -
Mitoura (seven species), Cyanophrys (one SP euee! and the
nominal subgenus Callophrys (six species). The six species
included in the nominal subgenus are characterized by green
on the undersides of the wings, with or without presence of
white maculations.

Callophrys (C.) afftnis (Edwards) has a light green
color usually without any maculations. This species ranges
from northern Utah and Wyoming to Washington and British
Columbia, and according to Clench the life history is still
unknown. Two subspecies C. a. affints (Edw.) and C. a.
washingtonta Clench currently are recognized, separable on
the basis of scale color dorsally and ventrally (Tilden,

1963).

A related species, C. sheridani (Carpenter), is charac-
terized by a solid line of white on the ventral surface of
the hindwings. There are three subspecies, C. s. sheridani
(Carpenter), C. s. neoperplexa (Barnes & Benjamin), and C.
Ss. newcomert Clench, separable by the pattern and degree of
development of the macular band (Tilden, 1963). The latter
two are confined mostly to Washington and Oregon away from
the immediate coast, whereas the nominal subspecies occurs
in the Rocky Mountain states.

A third species, C. apama (Edw.) has two subspecies, C.
a. apama (Edw.) and C. a. homoperplexa Barnes & Benjamin,
the latter characterized by the absence of a macular band.
The nominal subspecies is characterized by the maculations
on the undersides of the hindwings tinged with black rather
than the dominant white seen in other species in the subgenus.

Callophrys comstockt Henne, a recently described species,
occurs in semi-arid to arid habitats from northern Inyo
County in California south to the Providence Mountains of
San Bernardino County. C. comstockt appears to be closely
hewated to C. agpama, but lacks the tricolored fulvous band
found on the undersides of C. apama (Henne, 1940).

The present study concerns two species found in Cali-
fornia, C. dumetorum (Boisduval) and C. viridis (Edwards).
The former was previously considered to include two subspe-
cies, C. d. dumetorum (Bdv.) and C. d. perplexa Barnes &

1 A recently described population from the higher eleva-
tions of the Sierra Nevada has been named C. lembertt by
Tilden (1963), but not enough evidence concerning life
history and distribution is known to confirm its specific
status and relationships.

h Introduction

Benjamin, occurring from British Columbia to Baja California
Norte, with perplexa said to be confined to San Diego and
Los Angeles counties, and adjacent portions of Baja Califor-
nia. The ventral sides of the hindwings of this species are
a light green with a variable number of white maculations.

A third entity was recently described from the Pacific
Northwest. as C. d. oregonensis, based on restriction of the
gray scales to the Cuz vein on. the undersides ,of the ypene-
wings as well as on comparative notes on this entity's bi-
ology (Gorell tek), 19700)

The last species, C. viridis (Edw.) which also has a
variable number of white maculations, is known only from San
Mateo County to Mendocino County, California, along the im-
medtate coast. The deep yellow to blue-green tinge separ-
ates C. ptridis superficially from C. shertdani and C.
dumetorum.

Barnes & Benjamin (1923), Clench (1944) and Tilden (1963)
in characterizing differences between species in the subgen-
us Callophrys, included morphological features such as scale
color (wing fringes, forewing costa below, shade of green
below, labial palpi, facial hairs, and dorsal ground color)
and shape or angle of forewing and hindwing margins. After
examining a large series of specimens of the two so-called
species, Callophrys dumetorum (Bdv.) and Callophrys viridis
(Edw.), the present author found no evidence which seemed
conclusive or sufficient to warrant separation of the two as
species. These characters vary considerably even within a
single population.

Populations, of, the two were found to be al lopatmmc yin]
dicating that interbreeding probably does not occur. In
the absence of evidence of interbreeding, biological charac-
teristics such as differences in female ovipositional sub-
strate and differences in courtship and mating are charac-
teristics which were examined in an attempt to confirm or
refute a specific level of differentiation between the two.
Comparisons of larval morphology, host specificity, and de-
velopment time were also used to help determine whether the
two are to be considered subspecies or species.

ACKNOWLEDGEMENTS

Thanks are due to Dr. Jerry A. Powell, University of
California, Berkeley, for assistance in the preparation and
format of this study, as well as in the reading of the manu-
script. Also helpful were Drs. Robert Ornduff and Howell
Daly both of the University of California, Berkeley, in read-
ing and criticizing the manuscript. | also wish to thank
Paul A. Opler, University of California, Berkeley, for his
advice concerning research techniques and choice of study
Sites. Special thanks also go to Oakley A. Shields j3tenmenty
of La Mesa, California, for data and observations concerning

Two species of California Callophrys 5

the bionomics and behavior of a southern California popula-
tion of Callophrys dumetorum; John Emmel, San Francisco,
California, for observations on a population of Callophrys
vtrtdis in San Francisco, and David McCorkle of Monmouth,
Oregon, for life history data of C. dumetorum in Oregon and
Washington.

Grateful thanks are also due to the following individu-
als who allowed the examination of specimens from institu-
Piemalian private collections in theitr-care: Dr? Paul «H.
Arnaud, Jr., California Academy of Sciences, San Francisco;
Dantes. Bauer, Bijou, California; Dr. Ernst S. Dornfeld,
termvaiiis, Oregon; Dr. Stanley G. Jewett, Jr., Portland,
Oregon; Lloyd Martin, Los Angeles County Museum, California;
David L. Mays, University of Florida, Gainesville; David V.
McCorkle, Monmouth, Oregon; Dr. E.J. Newcomer, Yakima, Wash-
ington; Paul A. Opler, and J. A. Powell, University of Cal-
mromnta, Berkeley; R.0. Schuster, University’ of California,
Bayes: si. OO: E. Sette, Los Altos, California; Oakley Shields,
oumemhy. OF La’Mesa, California; Fred T.:Thornme, El Cajon,
California; Ronald S. Wielgus, Phoenix, Arizona.

iyesimcene- Ehanks go to Perry E. Turner, Jr., Paul A.
Opler and Alfred Blaker, all of the University of California,
Berkeley, for photographing the adults and young stages used
in the study, and to Dr. Kenneth Hagen, Division of Biologi-
cal Control, Albany, California, for identification of the
hymenopterous parasite reared on Callophrys virtdis. John
Emmel kindly contributed 28 pupae (20 C. dumetorum, 8 C.
viridis) and Dr. John Downey, University of Northern lowa,
Cedaimebatls, toaned several Callophrys pupal shells. Dr. R.
Bacigalupi, Jepson Herbarium, University of California, Ber-
keley, determined several plant species mentioned in the
TEX.

HISTORICAL REVIEW

Linnaeus, in 1758, proposed the genus Papilto which in-
cluded a diversity of butterflies. A single hairstreak
characterized by the complete green undersides of the wings
was named Papilio rubt. The name Callophrys was originally
propounded by Billberg (1820) to include three groups of ly-
caenids based on possession of three tails, two tails and
ene tail. ln the group characterized by one tail only P.
rubt was recorded. P. rubt was selected as the type of the
genus Callophrys by Scudder in 1875 although the more encom-
passing generic name Thecla remained in general usage until

1914.

Boisduval (1852) described Thecla dumetorum which he be-
lbreved to be a local race of the European species.

W. H. Edwards (1862) described Theela virtdts, charac-
terized by the uniform deep green wing undersides and by
white antennae.

6 Methods and Materials

Barnes & Benjamin (1923), after examining a small ser-
ies of Callophrys dumetorum from the San Diego area, named
the southern California race perplexa on the basis of com-
plete or almost complete absence of white maculations on the
undersides. They also believed Callophrys viridts to be
conspecific with the typical northern California race of
Callophrys dumetorum, rendering C. virtdts a synonym of C,
dumetorum.

Comstock (1927: plate 50) pictured a male and female of
C. viridis from San Francisco which he labeled '"'C, dumetor-
um, but C. dumetorum, in reality, does not occur in San
Francisco,

Clench (1944) revised the genus Callophrys and cited
C. vtrtdits as a species. San Francisco was named as the
type locality and a neotype was designated.

Ziegler (1960), in his revision of the Lycaenidae based
on male and female genitalia, combined Mttoura Scudder,
Sandta Clench and Ehrlich, and Callophrys Billberg under the
latter, making each a subgenus.

Tilden (1963) discussed many diagnostic characters use-
ful in separating the members of the subgenus Callophrys,
emphasizing scale patterns and wing shapes.

Gorelick (1970) described Callophrys dumetorum oregon-
ensts from the Pacific Northwest based on scale patterns and
biological data.

METHODS AND MATERIALS

The study was divided into four stages, the first being
the examination of 782 museum specimens (both species). Di-
agnostic characters were sought other than those having to
do with scale patterns in order to separate not only C. du-
metorum from C. viridis but also’C. d. dumetorum trom Guede.
perplexa. Dissections of male and female genitalia from
both species were done using the technique outlined in
Ehrlich and Ehrlich (1961), these specimens being chosen
from many different localities for possible variation between
populations.

The second stage dealt with obtaining specimens (inclu-
ded in the study) from many lepidopterists to clarify the
geographic range of these species. Weekend trips were taken
during the late spring months of 1967 to extend the range of
Lt vVErvadst,

The third dealt with the selection of several locali-
ties in the San Francisco Bay area where ecological and be-
havioral data for the two species could be obtained and com-
pared during the spring of 1967. Study sites were the
following:

Two species of California Callophrys 7

Callophrys vtridis:

1. San Bruno Mountains, near Daly City, San Mateo Co.
2. 9 mi. NE of Pt. Reyes lighthouse, Marin Co.
3. Bele Puerto Canyon, 22 mi. W. Patterson, Stanislaus Co.

Callophrys dumetorum:

1. San Gabriel Canyon, near Azusa, Los Angeles Co.
foeasane dunes, 1/2 mt. E. of Antioch, Contra €osta Co.
3. Brannan Id. State Park, Sacramento Co.

Observations of C. dumetorum were also made at Diction-
ary Hill (San Diego Co.) by Oakley Shields (via correspon-
dence). Specific localities within each study site were
chosen for capturing, marking and releasing adults in order
to study individual behavior.

The San Bruno Mountains population was observed from
February 10 to May 20, whereas Antioch and Brannan Island
were visited from April 4 to May 30. Observations were made
at the Pt. Reyes study site between April 30 and June 3, and
Del Puerto Canyon was only visited twice, in late March and
in mid-May. Field observations were made two or three times
each week at all but the latter two localities.

Adults of both species occurred at specific hilitop lo-
calities in the San Bruno Mountains (C. viridis) and Brannan
Island (C. dumetorum). Marks were made with a black felt
pen on the green scales on the wing undersides while the
Specimen was held within the net. The marks were made so
that the specimen could be determined as to date of marking.
To avoid loss of individuals immediately after marking, they
were held in the net until docile and then released. Mark-
ing was done all day for one day at each of the above local-
ities and all recaptured specimens taken throughout the
study were recorded as to date and time of day taken. No
secondary marking or re-releasing of the recaptured speci-
mens was conducted.

Adults were observed in the lab on several plants in-
cluding the natural host plants but no courtship, mating or
oviposition occurred in the lab.

The fourth stage included the lab rearing of larvae of
both species on a series of related and unrelated plants to
determine relative specificities and also to note the rate
of development on each. The experimental host plants used
were:

1. Lupinus arboreus Sims. (Leguminosae)

2. Vicia sp. (Leguminosae)

3. Lotus scoparius Ottley (Leguminosae)

4. Trifolium obtusiflorum Hook. (Leguminosae)

5. Cytisus monpessulanus L. (Leguminosae)

6. Eriophyllum staechadifolium Lag. (Composi tae)

8 Geographic and Ecological Distribution

7. Gnaphaltium sp. (Composi tae)

8. Achillea lanulosa Nutt. (Compositae)

9. Amsinekta tntermedia F.& M. (Boraginaceae)

0. Ertogonum fasciculatum Benth. (Polygonaceae)

1. Ertogonum lattfoltum lattfolium Sm. (Polygonaceae)

Eggs of C. dumetorum were obtained at the Antioch and
Brannan Island study sites by taking large numbers of Lotus
scoparius buds from the field to the lab for microscopic ex-
amination. C. vtridts eggs were taken from the leaves of
Ertogonum ltattfottum Latitfotzum at Pt... Reyes.

Glass vials were used, one individual per vial, to rear
larvae of both species. Each vial was labeled according to
date of egg hatching and arranged by locality in a rack for
daily observations. Larval development during the 1967
season was not recorded in detail, but successive instars
were accurately recorded in the 1968 lab data using head
capsules as an instar number indicator. One or two leaves
and several flowers, were placed in each vial with the egg
or larvaand replaced every other day when desiccation or
wilting occurred. Records were kept of the larval accept-
ance of each plant, especially the time taken to feed, the
manner and/or duration of feeding, and of larval refusal to
feed. The rate of development on each plant was used as an
index for feeding success as well as host acceptance. Lab
temperatures during the rearings (1967 and 1968) ranged from
AZe stom Pore ke

Developing larvae were examined under 45x for distin-
guishing characteristics and an ocular micrometer was util-
ized at 32x for larval head capsule measurements. Life his-
tory photographs were taken (Figures 1-14) and comparative
morphology of the immatures were diagrammed (Figures 15-21B).
Larvae were fixed in KAAD for 20 minutes and preserved in
95% ethyl alcohol. Many of these larvae, along with all
marked and recaptured specimens have been deposited in the
California Insect Survey collection at the University of
California, Berkeley.

GEOGRAPHIC AND ECOLOGICAL DISTRIBUTION

Callophrys dumetorum (Bdv.) occupies a wide distribu-
tion extending from British Columbia to Baja California Nor-
te, eastward to the eastern edge of the Sierra Nevada in
California. Sierra Nevada populations show broad ecological
tolerance, occurring as high as 5000'. Callophrys viridis
(Edw.) is found along the immediate coast in California from
the San Bruno Mountains in San Mateo County to Juan Creek,
near Westport, Mendocino County (see map 1). Efforts to
find this species along the coast in Oregon and the northern-
most boundaries of Humboldt County failed. Neither C. dume-
torum nor C. vtridts occur in Lower Sonoran areas.

OO ——————

Two species of California Callophrys 9

42)

DISTRIBUTION OF CALLOPHRYS DUMETORUM (Bdy.)
IN NO. AMERICA

38
37

37)

4) CALLOPHRYS VIRIDIS (Edw.)
) CALLOPHRYS DUMETORUM (Bdy.)

HB VARIANT CALLOPHRYS POPULATION,

CALIFORNIA INSECT SURVEY
Department of Entomology and Parasitology

UNIVERSITY OF CALIFORNIA ° BW

25 50 : > = : 2 Rak
ORAFT 1955 Waser
15 i“

istribution of Callophrys dumetorum and C. viridis
n California.

PAPEL. ivy D

10 Comparative Morphology

FLIGHT PERIODS

Populations of C. dumetorum fly early in February in
southern California whereas adults in San Francisco Bay area
populations are not seen until mid-March. At Kusshi Creek,
Yakima County, Washington, flight does not occur until May
(Newcomer, 1965). The C. viridis population studied in the
San Bruno Mountains reached a peak flight period on March 7,
1967 whereas the C. vitrtdts population found along the sand
dunes at Pt. Reyes on the coast in Marin County did not
reach a peak flight period until May 14, 1967.

Host ASSOCIATIONS

Ertogonum lattfoltum latifolium Sm., the observed food-
plant of C. viridis (Brown and Opler, 1967) is only known
from the coast of California and Oregon, whereas C, dume-
torum females have been observed ovipositing on species of
Syrmattum, Hosackia, Lotus and Eriogonum in southern Califor-
nia (Coolidge, 1924). Newcomer (1965) lists ©. dumetorum
foodplants in Washington as Eritogonum heracleotdes, E. com-
posttum and £. elatum. The hostplant of a Varteanhtepepeia] |
tion found at China Flat Campground in the Sierra Nevada is
probably Eritogonum wrtghttt Torr. ssp. traechygonum (heres)
Stokes whereas the foodplant of the Del Puerto Canyon popu-
lation, on the basis of three Callophrys eggs found on a
single stand in March, 1968, is Eriogonum Latifoltum Sm.
ssp. aurtculatum (Benth.) Stokes.

COMPARATIVE MORPHOLOGY
EVALUATION OF CHARACTERS: CALLOPHRYS DUMETORUM AND C, VIRIDIS

Genital dissections were carried out using specimens ta-
ken in two localities for each species. No differences were
discovered between the males of C. viridts and those of C.
dumetorum. The male gnathos and valvae appeared to have the
same length and the same degree of sclerotization. The fe-
males of both species, too, had no structural differences.
Careful examinations of sternites VIII, IX and X yielded no :
differences in size or shape. The signum was examined for
possible differences but the amount of sclerotization was
great in worn specimens while barely present in fresh ones,
evidently a function of age. Male and female genitalia of |
C. lembertt Tilden and CC. comstockt Henne were also examined :
but showed no apparent structural differences from one ano- :
ther. Therefore these structures do not provide the answer
in determining the validity of species.

Androconial scales from the scent pouches of the males
of both species were removed with a minute probe and placed
on a slide for examination under 100x. No apparent differ-
ences were seen. Labial palpi from both males and females
of both species also manifested no structural or scale differ-
ences. Of the fourteen characters mentioned by Tilden (1963)

Two species of California Callophrys 1]

for separating the members of the subgenus, ten were insuf-
Ficient after examination of 150 specimens of both species.
Such a character, for example, is the number of white macu-
lations present on the undersides of the wings mentioned by
Barnes & Benjamin (1923), Clench (1944), and Tilden (1963)
as a good diagnostic character. But according to Ford
(1945), individuals within a single population vary in the
number of maculations present and do so even in the Euro-
pean species.

Examination of 596 specimens of C. dumetorum and 186 of
C. viridis (total of 782 specimens) yielded the following
characters useful in separating the two species:

1. Females of C. viridis are a uniform olive-gray on the
damsal surface, as are the males (fig. 7), varying to
a mild olive-brown suffusion in some specimens. C,
dumetorum females are dark gray-brown with a variable
amount of yellow-brown suffusion within a single popu-
lation as well as throughout the entire flight range

fhegs. 0, 9).

2. The color of the ventral surface of the wings appears
to be a soft or dull yellow-green, varying to olive-
green in almost all California populations of C., du-
metorum (figs. 10-12). C. viridis, on the other hand,
has deep yellow to blue-green undersides varying only
with occasional mild yellow-green individuals within
a single population as well as throughout the entire
range (figs. 13, 14)

3. The antennae, when viewed dorsally, are black and
white in C. dumetorum whereas fresh C. viridis speci-
mens have all white-scaled antennae.

4h, In most California populations of C. dumetorum, the
green scales on the ventral surfaces of the forewings
medially never exceed the Cu] vein whereas C. viridis
populations examined almost always have green scales
reaching the Cuj or Cug vein of the forewings.

Gharacters |] and 2 are evidently the most efficient way
to separate C. viridis from C. dumetorum. The third charac-
teristic is useful only when the specimens being compared
are fresh. C. virtdts antennae tend to lose the white annu-
lar scales after a few days of flight, making them appear
similar to those of C. dumetorum.

Of 68 randomly selected C. dumetorum specimens (both
male and female) examined from all over California, 4 (5.8%)
had the medial green scales on the forewing undersides be-
tween the costal and radial veins, 17 (25.0%) had green
scales extension restricted to the M; vein, 32 (47.1%) had
green scales reaching the M9, 12 (17.6%) had green scales
reaching the M3 and 3 (4 4%f showed green scales reaching
Ee Cu |.

12 Comparative Morphology

On the other hand, 75 examined C. vtrtdts specimens
from Pt. Reyes and the San Bruno Mountains showed 5 (6.6%)
with green scales restricted to the Mo, 68 (90.1%) with
green scales reaching Cuz sand 2 (3.333 with green reaching
Cu9.

EVALUATION OF CHARACTERS: CALLOPHRYS DUMETORUM PERPLEXA

Other questions raised concern the validity of Callo-
phrys dumetorum perplexa, the atypical race found in south-
ern California. Barnes & Benjamin (1923) recognized this
race as a valid one occurring in Los Angeles and San Diego
counties southward into Baja California Norte and distin-
guishable from typical C. dumetorum by ''the complete or al-
most complete absence of white markings on the undersides".

Sixty two specimens .of Cend. perplexa were examined
from the vicinity of the type locality (San Diego) yaemor
which were males. These males included 6 individuals with
no white maculations on the wing undersides (11.3%), 8 with
1 (15.1%), 36 with 2 (67.82) and. 3 specimens hain)

Of the 9 females examined none lacked any maculations,
2 had only 4. (22.2%), 2 had 2(22.2%), and 3 showed mene)
The remaining 2 females had more than 3 (22.2%) (see graph
1A).

In comparison, a series of 51 males of C. dumetorum ex-
amined from Brannan Island State Park, Sacramento County,
varied from none without any maculations, 3 with 1 (5.9%),
© Ae 2 (68.6%), 8 with 3 (15.7%), and 5 with: meme meiner

9.8%).

Females of the above population also varied considerably.
Of the 21 specimens examined, 2 had no maculati ons oaacee
2 had J (9.5%), 8 had 2 (38.1%), 6 had) 3 (28.6%) .:anideeeeeten
ed more than 3 (14.3%) (see graph 1B).

The resuits of the above comparisons suggest that north-
ern California populations are characterized by more spots
but not significantly enough to warrant subspecific recogni-
tion. Because the males (and most females) of both popula-
tions compared showed that the two-spot condition was most
prevalent, clinal variation was not assumed to be the case.
These maculations, although variable in number, never vary
in scale pattern. All are white outwardly and black inwardly. |

The invasion of the brown scales on the undersides of
the forewings almost to the costa, thus restricting the green
Scales to the costal margin area, is a character which is
supposed, to. be; typically on. Gnd. perplexa. No differences
could be found, however, concerning the restrictions of green
scales in any of the C. dumetorum examined throughout its en-
tire range in California.

Two species of California Callophrys 2)

GRAPH 1 - A comparison of maculation percentages between two
California populations of Callophrys dumetorum:
A,- Southern California population. B - Northern
California population.

14 General Biology

The color of the scales composing the fringes of the
wings was used as a character by Barnes & Benjamin (1923)
separating the perplexa form from the typical form of C. du-
metorum. These fringes were dark and becoming white outward-
ly, seen in typical C. dumetorum as well as in C. d. per-
plexa. This, too, is a variable trait of no 'stqmit peamees

Lastly, Clench (1944) asserted that C. d. perplexa is
characterized by the "presence of fulvous edging (light
brown or dull yellow) on the costa of the forewings ventral-
ly". This ts.a characteristic also seen in the nortnermn
California specimens of C. dumetorum.

After review of 554 specimens relative to the above
four characters attributed to Callophrys dumetorum perplexa,
this writer concludes that they do not validate a distinct
southern California subspecies because of their presence in
the phenotypes of northern California populations as well.
Because of the population variation factor present in Cali-
fornia, no nomenclatorial distinction is required.

The chief enigma encountered during this study was the
discovery of two populations of Callophrys (s. str.) that
were not clearly either C. dumetorum or C. viridis. A female
Callophrys examined from China Flats Campground, El Dorado
County, appears to be C. virtdts although populations repre-
senting the C. viridis phenotype are not otherwise known to
occur in the Sierra Nevada. A second interesting population
was found in Del Puerto Canyon in western Stanislaus County.
Here individuals match the original description of C. virti-
dts with the exception of the black annular scales present
on the antennae which fit C. dumetorum. In several specimens
from Del Puerto Canyon, the green scales were restricted to
the area above the subcostal veins as seen in C. dumetorum.
Since these characters were seen even in fresh specimens,
the question must be raised as to whether C., dumetorum and
C. viridis interbreed. Field work throughout the Bay Area
where the two come closest together while remaining allo-
patric indicate that they do not.

In summary, the failure of good evidence of interbreed-
ing suggests the two are reproductively isolated species,
and the failure of comparative morphology to reveal reliable
criteria for separation of the two indicates other means of
confirming their distinctness must be sought. Biological
characteristics are therefore examined in this study.

GENERAL BIOLOGY

Most of the species in the subgenus Callophrys are sin-
gle brooded, fly only during the spring and feed on a fairly
wide range of hostplants. Females of the Palearctic species
Callophrys rubt (L.) oviposit on rock-rose (Cistaceae),
purging buckthorn (Rhamnaceae), gorse, broom, Dyer's green-
weed (Leguminosae), bramble (Rosaceae, dogwood (Cornaceae)

—<—— —— _—-—— + or

i

Two species of California Callophrys 15

and whortleberry (Ericaceae) (Ford, 1945). Most Nearctic
species use members of the buckwheat genus Eriogonum, with
C. dumetorum also using the leaves and buds of legumes such
as Lotus (some genera recorded in older literature as food-
plants for C. dumetorum, i.e., Lotus, Hosackia and Syrma-
titum, recently have been included in the genus Lotus (Munz,
1963)). Larvae feed on the leaves and flowers and appear to
derive protection from predators due to their cryptic color-
ation. Pupation occurs in inconspicuous places at the base
of the hostplant (Comstock, 1927).

General adult activity is influenced by weather condi-
tions. Clouds blocking the sun, temperatures below 55° F,
wind and rain prevent flight activity. During cold weather,
adults lean or flatten themselves against leaves, rocks and
other flat surfaces. Presumably this is a thermoregulatory
mechanism through increased area for insulation (Clench,
1966). On warmer, sunny days flight activity is seen all day.

BIOLOGY OF CALLOPHRYS DUMETORUM

The biology of C. dumetorum was studied at four locali-
ties, the first being San Gabrie] Canyon (near Azusa), Los
Angeles County, an area characterized by many chaparral spe-
cies, including Ertogonum fasctculatum and Lotus scopartus.
ftuerese cond locality was Dictionary Hill, San Diego County,
where observations were made by Oakley Shields in early
March for several weeks. The third locality, 1 1/2 miles
Edsmean Anttoch in Contra Costa County, is characterized by
a river delta sand dune flora. The last locality, Brannan
Island State Park in Sacramento County is also typified by
a sand dune flora. Observations were made on seven days be-~
tween April 8 and May 11 at the latter two localities.

ApULT BEHAVIOR

Callophrys dumetorum adults, flying not more than three
or four feet above the ground, alighted repeatedly on both
Lotus seoparius and Eriogonum fasciculatum in San Gabriel
Canyon. At Antioch, however, females circled specific stands
of Lotus and landed frequently, ignoring the numerous stands
of Ertogonum latifolium ssp. present. This lack of interest
in E. Latifolium by C. dumetorum females was seen throughout
the 1967 flight season and indicates that this plant is prob-
ably not used as a host. "Aggressive'' behavior was display-
ed by many males, especially when another butterly species
entered the area. The possibility, however, that these
flights were merely an attempt to elicit female response
should not be ruled out. Not infrequently two C,. dumetorum
males perched within one or two inches of one another and
numerous "scuffle flights'' or mutual response reactions
(whereby males compete for a given territory while in flight)
occurred, those individuals involved revolving around one an-
other up to 20 feet in the air, returning to separate leaves

i Callophrys dumetorum

on the same plant or to different plants. Such behavior has
been seen in many butterly species and is discussed in de-
tail by Shields (1968).

At Brannan Island 10 males and 9 females were marked
and released at 10:00 A.M. on April 13, but of they tOage-
males and 42 males taken between April 13 and May 2, only 1
marked male was recaptured.

In March 1967, Oakley Shields marked individuals of ¢,
dumetorum for 18 days to determine whether males attempt to
occupy specific territories. The markings occurred @npic—
tionary Hill in San Diego County, an area characterized by
an abundance of Brassica nigra. Of 20 males marked, 7 (35%)
were recaptured as follows: 2 after 1 day, 1 after 4 days,

l after 5 days, 1 after 8 days, 1 after 9 days, and 1 after
18 days. The worn male taken after 18 days was not in ex-
cellent condition when marked, giving some indication of the
longevity of individuals of this species.

Courtship behavior, observed several times at Brannan
Island during the 1967 season, commenced as early as 10:15
A.M. (PST) when an individual male approached a female in
flight for several seconds, ''forcing'' the female to alight.
The female, facing the shrub, perched motionless while the
male perched on a leaf behind the one occupied by the female
and vigorously vibrated its wings. It then moved in a clock-
wise direction 180° until facing outward in the opposite
direction. The entire courtship process lasted no longer
than 15 seconds.

As a summary to both Shields' and the present author's
observations, mating occurred on the summit in a given terri-
tory assumed by a single male. The territories assumed by
males of C. dumetorum during these observations (as shown by |
the marking study results) seemed to shift around when many
males were present.

Mating behavior was witnessed at 12:30 P.M. (PST) at
Brannan Island on April 18, 1967 when one pair was observed
for 30 minutes. The female faced the interior of the shrub
(Lupinus arboreus) with the antennae oriented upwards whereas
the male faced the opposite direction, antennae pointing
downward. Later, another pair was observed in copulo on Lu-
pinus from 10:56 A.M. to 11:45 A.M... In-April, 1966anenmeiie
mating pair was observed displaying the above posture as late
as 3:00 P.M. on Lotus. scopartus. After several observed ma-
tings, lab dissections of four females (two of which had
been observed mating) were carried out. The two mated females
yielded two concentrically-deposited spermataphores in the
mouth of the bursa copulatrix indicating that many females
mate at least twice.

Oviposition was seen at 10:00 A.M. on May 5, 1967 when a
female, perched on a Lotus bud, inserted its abdomen well
Into the bud and deposited a single green egg 90 seconds after

Two species of California Callophrys 17

insertion. After turning 180° to face the interior of the
same bush, the butterfly flew to another apical bud and ovi-
posited another egg after only 30 seconds. Rapid wing rub-
bing ensued, followed by antennal vibration. Only when the
antennae touched another bud or leaf surface did the female
move onto another part of the plant. The female inspected
many buds and leaves before attempting to oviposit and flew
off if a satisfactory substrate was not found. At no time
did a female C. dumetorum at Brannan Island attempt to ovi-
posit on Ertogonum latifoltum ssp. No eggs or larvae were
collected from Ertogonum fasetculatum in San Gabriel Canyon
thus failing to support Coolidge's claim that females accept
Ene Vatter as a suitable host. Since #. fasciculatum is
often found in association with Lotus seoparius throughout
the California coast ranges, Ertogonum should not be ruled
out as a foodplant for some populations.

LIFE HISTORY

The eggs, approximately 0.4 mm in diameter, are ovoid,
Finely reticulated, green, and are deposited singly on the
host plant (figs 1, 15). Eggs were collected from apical
buds of Lotus scopartus at Antioch and Brannan Island and
kept at room temperature. Emerging larvae were placed on
experimental foodplants, and the results of these rearings
are presented in Tables 1 and 2.

Morphological changes in each instar in the Brannan
Island population corresponded closely to the descriptions
given by Coolidge (1924). He showed Callophrys dumetorum to
have four larval instars, verified by measurement of the
head capsule widths of each larva examined. This technique
has been found to be useful; there is apparently no overlap
of head capsule width between instars in C. dumetorum and
C. viridis larvae. This may be true of most lycaenids since
it was also reported by Lawrence and Downey (1967) in lar-
vae of a plebejine lycaenid, Everes comyntas Godart.

The following measurements were made on a total of 23
C. dumetorum larvae reared and preserved during 1967 and 1968.

Head Capsule Width (mm) Body Length (mm)
l instar: (N=7) 0.15-0.25 0...92='1. 54 stave. 1.172
(avg. 0.20)
I) instar: (N=1) 0.40 S20
Pel-instar: “(N=4) 0.70-0.92 4.80-9.10 (avg. 7.59)

(avg. 0.80)

IVoinstar: (N=11) 1.23 11.8-15.5 (avg. 14.8)
Cave 1223)

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18

Two species of California Callophrys

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20 Callophrys dumetorum

TABLE 2 - DEVELOPMENT OF CALLOPHRYS DUMETORUM LARVAE ON
LoTUS SCOPARIUS LEAVES (1968)1

Egg 1st 2nd 3rd 4th Color Pipa
Insrar “anstar 2nSstar yy. tisiade changes
IV-7 IV-7 IV-11 IV-16 IV-21 Pink IV-30
IV-26
IV-7 Iv-8 IV-11 IV-16 IV-21 Pink IV-30
IV-26
IV-7 IV-8 IV-12 IV-19? IV-25 Pink V-3
IV-29
IV-7 IvV-8 IV-11 IV-21 IV-25 Pink V-3
IV-29

1 Eggs collected April 7, 1968, at Brannan Island State
Park, Sacramento County, California on Lotus secopartus.

Measurement of the molted head capsule widths at the
end of each instar yielded the following ranges:

| instar: 0.15-0.38 mm
ll instar: 0.39-0.68 mm
lll instar: 0.69-1.10 mm
(Vo Dnisttiaws 14 1= 03237 mm

The mature 4th instar larvae of C. dumetorum are uni-
form light yellow-green with subdorsal and lateral white
stripes, turning pink just before pupation (figs. 2asemeue
Triordinal crotchets arranged in a mesoseries are present on
the prolegs (figs. 18A, B) and the crotchets (fig. bones
interrupted at the middle by a spatulate lobe, stated by
Fracker (1915) to be typical of lycaenid larvae in general.
The body, covered by numerous secondary setae, has uniformly
rounded segmentation throughout the entire length. The head
capsule is heavily sclerotized from the first to the third
instars but only the ventral portion of the adfrontal sutures
(above antennae) and the ocellar area remain heavily sclero-
tized in the, fourth) ns tan Gnicemiy

Upon hatching, first instar larvae began to skeletonize
terminal leaves of Lotus scoparius; the second instars did
the same. The larvae moved down the stem when the upper
leaves died or became completely skeletonized. Mature larvae
defoliated the lower branchlets rather than skeletonizing
leaves. Larval feeding in general occurred throughout the
morning hours in all instars, somewhat actively in the after-
noon, tapering off to almost no feeding at all by early even-
Ing. Resting larvae positioned themselves on the stem or on
the tops of the leaves. Coolidge (1924) stated that the de-

Two species of California Callophrys 2)

velopment time from egg to pupa in a southern California pop-
ulation was 32 days with larvae feeding on Hosackia (Lotus).
Rearing results from the Brannan Island population indicated
a 30 to 35 day span on Zotus. Although feeding occurred
successfully on the flowers of Trt folium obtusiflorum and
Ertogonum lattfoltum latifoltum, development from egg to pu-
pa took as long as 39 days. Feeding on Ertogonum fascicula-
tum leaves resulted in one ¢, gymetorum larva reaching the
pupa stage in 36 days.

Unfortunately, a search for field larvae failed and all
life history work was carried out in the lab. Thus no para-
sitism was encountered during the course of the larval rear-
ings and only one mature larva was lost due to bacterial
disease,

The pupae (figs. 3, 20, 21B) are a uniform dark or
brown-black with the surface texture of the wing pads ex-
cessively sculptured, almost forming rows, in contrast to
the dorsal surface. A total of 26 were measured, with the
length ranging 8.6-10.0 mm (avg. = 9.27mm).

BIOLOGY OF CALLOPHRYS VIRIDIS

Observations on C, viridis (reported as "Theela dumetor-
um'') were made in the western part of San Francisco by Will-
iams (1910) who first determined the larval hostplant to be
Ertogonum latifolium latifolium. In June, 1963 J. A. Powell

tn litt.) collected ¢, viridis larvae in the San Bruno
Mountains in flower heads of Ff, 12. latifoltum and noted that
their pale color with pinkish markings blended quite well
with the flowers. Adults emerged in February and March the
following season. Further observations on C. viridis, em-
phasizing oviposition behavior, were made by Brown and Opler
(1967) iti the San Bruno Mountains.

C. viridis was studied extensively during the present
investigation at two localities in central California. The
first is the San Bruno Mountains near Daly City, San Mateo
County. This study site, located about 4 miles from the
Pacific Ocean, is characterized by steep rocky slopes with
dense, diverse coastal chaparral including an abundance of
BE. Lt. tattfolium. No Lotus secopartus was seen here. Obser-
vations were made on 14 days between March 2 and May 12, 1967.

The second study site, nine miles northeast of the Point
Reyes lighthouse, Marin County, consists of beach sand dunes
and associated flora, including £. LZ. lattfoltum. Like the
San Bruno Mountains, no Lotus seoparius was found. Cold
weather with low clouds and fog was prevalent during the
early morning hours (before 11:00 A.M. PST) but most days
warmed up considerably by noon when most activity was obser-
ved. Observations were made on five days between April 30
and May 28, 1967.

22 Callophrys viridis

ADULT BEHAVIOR

As in the case of most Callophrys species; Ci y7veaes
activity is greatly influenced by weather. Flight habits
and sunning behavior are similar to those seen in C, dume-
torum. Observations during the 1967 season were made with
difficulty at the San Bruno Mountains study site due to wind
and long periods of rain.

The courtship pattern exhibited by C. viridis is like
that of C. dumetorum; a female entering a given male terri-
tory was courted by one or more males while in flight. But
unlike C. dumetorum, males of C. viridis in the San Bruno
Mountains (due to their lesser numbers) were able to main-
tain a specific territory with a minimum of shifting around.
Two females and two males were marked in late March, 1967,
and two males were recaptured on the same plants five days
later.

Courtship, mating and oviposition behavior of Callophrys
vpirtdts was observed at Pt. Reyes. The first two behavioral
mechanisms were found to be similar to that observed in C.,
dumetorum at Brannan Island State Park. Oviposition was
witnessed on May 14, 1967, at Pt. Reyes at 10:00 A.M.; a
single female landed on many stands of Ertogonum, tapping
several leaves on each with the antennae. Finally, after
twenty minutes of flying from plant to plant, the female
alighted on an Ertogonum leaf; and, after touching the under-
side of the leaf with the antennae, bent the abdomen under-
neath and deposited a single green egg near the apical end
of the leaf after an estimated 15 seconds. The pilose mat
on the leaf underside made it almost impossible to perceive
the egg (photo 4).

Host selection by Callophrys virtdts females was limi-
ted at both localities to Ertogonum lattfolium . This may
be due in part to the absence of suitable leguminous plants
such as Lotus secopartus. Vieta sp. occurs at the San erunoe
Mountains site, but was found to be unacceptable to C. viri-
dis larvae in the lab (see Table 3).

John Emmel (in 1ttt.) observed a female C. viridis ovi-
posit on Lotus secopartus- at 1V23507ALN. (PST) in the Sunset
Heights district of San Franciseo, April’ 27, 1960. ."nneecee
was laid within the apical bud as observed during my study
of C. dumetorum females. The Sunset Heights locality is
characterized by an abundance of £. Llatifolium latifoltum,
and Emmel reared larvae taken from both plants between June
3 and June 10, 1968 at this site’ (west of the UsC: Medea
Center). According to Emmel, the San Francisco Presidio dunes
north of the Veteran's Administration Hospital support both
Lotus and Ertogonum and both are accepted by C. viridis
females.

No Ertogonum in the Pt. Reyes vicinity was yet in bloom
while the females were ovipositing. Oviposition on stands

Two species of California Callophrys 23

of FE. Ll. lattfottum which have reached the bloom stage, how-
ever, has been recorded by Brown and Opler (1967) in the San
Bruno Mountains. In this case the female walked slowly up
and down the inflorescence stalk of the hostplant before de-
positing the eqg on a leaf underside. It appears that no
matter where the egg is deposited the females of both species
of Callophrys display numerous inspection flights before the
hostplant is chosen.

LiFe HIsToRY

The eggs are identical in size, shape and color to
those of Callophrys dumetorum but on Ertogonum are deposited
singly on the undersides of leaves near the base of the plant
(fig. 4). Both eggs and larvae were collected from the
leaves of £. Ll. Latifolitum at Pt. Reyes in May of 1967 and
1968 and young larvae were offered the same series of plants
used to test host specificity in C. dumetorum. The results
are incomplete, however, due to disease and parasitism en-
countered during the rearings. The larvae in many cases
died before pupation. The results of the 1967 and 1968 rear-
ings are given in Tables 3 and

Body lengths and head capsule widths were measured in
38 C. viridis larvae as follows:

Head Capsule Width (mm) Body Length (mm)
! instar: (N=10) 0.31-0.38 1.4-2.9 (Avg. 2.225)
(Avg. 0.34)
1! instar: (N=11) 0.40-0.62 3.7-5.5 (Avg. 5.1)
(Avg. 0.57)
lil instar: (N=7) 0.69-0.99 4.9-9.5 (Avg. 7.0)
(Avg. 0.83)
IV instar: (N=10) 1.15-1.30 9.8-16.0 (Avg. 12.8)

(Avg. 1.25)

Measurements of the molted head capsule widths yielded
the same ranges found for C. dumetorum larvae, with possible
exception of the upper limit of the IV instar head capsule
width.

The first instar laryae examined were light green with
setation much like that described by Coolidge (1924) for C.
dumetorum. Red spots appeared along the subdorsal ridges
during the late third instar and remained in the mature fourth
instar, corresponding to the description given by Williams
(1910). Mature larvae were variable in color; some were
light yellow-green with a small amount of pink varying
slightly to red, whereas others, even within a single popula-
tion were whitish marked predominantly with red (figs. 5, 19).
Unlike the uniformly rounded segmentation in C, dumetorum,
larvae of C. viridis have pronounced dorsolateral and segment-
al ridges. The setation, crotchet arrangement and head cap-

Callophrys vtridts

24

(OT-IA :PeTtd)
a” EX

#(€-IA :PeTtd)
ge A

x(E-IA ?PeTd)

uohuey
oqjteng Ted

SOACST WNnZD7
-norosof wnuoborig

SIOMOTJ wnaio7zs
-zsnzqo unr, 0f 14a],

SIOMOTJ wnto zs

Bc7A mM -zsn4qo wn170f14 J
(peztTatsered

peTtd) SOACST wn170f1407
L-A QZ-A . wntz7of1,07 wnuobo1raig
(OIT-IA Petd) SuUTeqUNOW SoAeeT wn2170f[1401
G=TA Q2-A oundlg ueg wn17O0Of1707] wnuocbo1atg
GI-IA *petd Qc-A "09 OUTOOPUSW SOACOT
yoorg ueng snizofi~ssDpdo snz.0o7

peztatseized
(Cedi SSG) SOARST
EIUIN eX ii sniapdoas snq,o7q

peztatsesred
Cea = Pera) yYIeg o4e49 SOACST
(GN Bc-A “ST ueuuedg sniapdoas snz,07

ICY4SUT ICS1SUT HEASOME I@YSUT KAREZOO TE
edngd 44 pag pug AST 334 que, dq soy LNVIdLSOH

(/96T) 3VAYV] SIGINIA SAYHdOT1V) N3ILYNO4 4O LNAWdOTSASg GNV JONVLd3IOW LNVIg LSOH - ¢ 37aV]

25

Two species of California Callophrys

psss 04 4dweq4e ON,

(foe aI
-peTtd)
oN

(peztqtse
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BLA

OWN, EIS) rap)
ha A,

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-nssadsuow sns21zfg
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SOARST

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SIOMOTJ wn170f1407
unrqzof44y7 wnuoborg,

wuny1 of 2404
ountg ueg wnrqzo0of1407 wnuoborag

uokueg SOACST unyD7
O4teng [eq -norvospnf wnuoborary

26 Conclusions

sule morphology are identical to those in C. dumetorum. The
distance between the lateral adfrontal sutures proximal to
the clypeus (widest point) were identical for mature larvae
of both. species. {avg. 0). 70 mm)’.

An undescribed species of Apanteles (Hymenoptera: Bra-
conidae) was recovered from one second, two third and one
fourth instar larvae during the 1967 rearings. Thompson
(1944) recorded no parasite record for any species of Nearc-
tic Callophrys, but several ichneumonid and tachinid species
are known to parasitize C, rubt in Europe.

Pupae were examined from Pt. Reyes and the San Bruno
Mountains and their lengths are: (N=14) 8.2-9.7 mm (avg.
9.2 mm)?

These pupae, unlike those of C. dumetorum described
above, are brown with a mottled appearance owing to inter-
spersed pale areas, particularly on the wing covers. Both
species exhibited finely sculptured ridges, but these appear
less pronounced on the lighter colored pupa of C. viridis
(figs al 6. | alka

RESULTS AND CONCLUSIONS

Laboratory and field study during the spring of 1967
and 1968 on Callophrys viridis and C. dumetorum produced
interesting similarities between the two. Host selection
(based on host availability) in the field proved to be of
little use in separation of the two species, while larval
characteristics provided the most valuable diagnostic infor-
mation. The results of the study are summarized as follows:

1. Laboratory inspection of 63 San Diego County Callo-
phrys dumetorum specimens showed that the four characters
used most frequently to define the so-called subspecres me.
d. perplexa were found to be present in northern California
populations in varying proportions. The degree of ditftern-
ence does not appear to warrant treatment of C. dG. perpvema
as a valid subspecies.

2. The percent difference in green scaling on the wing
undersides between C, viridis and C. dumetorum is consistent
enough to distinguish between them; ground color on the fe-
male uppersides and the quality of green color on the under-
sides are also useful in separating the two entities.

3. At no place in the respective distributions of the
two species has interbreeding been found to occur, although
allopatric San Francisco Bay area populations of both species
fly during the same months.

4, The number of individuals marked and released for
each species was too small to be of statistical significance,
but the number of recaptures along with observations of male

zy

Two species of California Callophrys

ee- TA

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(yutd)
Gple A

(murd)
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untzzof1407 wnuoborag

SoAvST wn170f1407
unz70f$1407] wnuobo1dgy

SOAPST uwntz0f1407
un?t.,of14z071 wnuoboragy

SOACST wntz70f{1407
untz,0f[14071 wnuobo1aty

SOAVOT wn170f1407
untzof414zp7] wnuobortg

a 8 eS eh ee

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HOLoD

Asse OLA: SenA u
YIeI 3484S
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(PeTd)
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(peTd)
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(PezT4Tsedeg)
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(PeZTITSe1ed)
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(8961) AVAYVT SIGIYIA SAYHdOTIV) 40 LNaWd013A3( JHL NO SNOILVAYNSSE) - 4 JTEV]

28 Coneluatons

concentration points indicate that courtship behavior for
both species is identical. Territories, while sustained by
individual males for short times, shift around from day to
day when numerical density is high, but appear to be main-
tained over several day periods in sparse colonies.

5. Aeriai courtship and reverse male-female orienta-
tion on the host and non-hostplant substrates were seen to
occur at least twice by individuals of both species.

6. Mating was observed twice during the morning hours
in both species; courtship behavior and oviposition also
were witnessed in morning and not commonly during afternoon.
Undisturbed males and females of both species remain in ecop-
wlo. for at Veas ti. 310" mi mies.

7. Females of C. dumetorum select Lotus secopartuyey for
oviposition even when Ertogonum latifoltum is present; while
C. vtrtdts females select Lotus and/or Ertogonum when both
are present in the same locality. Females of both species
spend much time examining and probing the substrate before
egg deposition, evidently in relation to the structural
quality of the hostplant.

8. In the laboratory, larvae of C. dumetorum fed suc-
cessfully on Lotus scopartus, Ertogonum lattifoltum, EF. fas-
etculatum and Trifoltum obtustflorum; C. vtrtdis larvae fed
successfully on the first three but failed to accept J7rt-
TOCeum.

9. Larvae of both C. dumetorum and C. vtrtdis matured
more rapidly on Lotus scopartus than on any other host.

10. Immatures provided the best morphological charac-
ters for the separation of these two species. Most note=
worthy are the characters given in Table 5.

As stated by Clench (1963), Callophrys is an inter-
esting subgenus because of the difficulty in discriminating
between the species comprising it. The results of the pres-
ent study indicate that the morphology of the larvae lend
strongest evidence for confirmation of species integrity in
C. dumetorum and C. viridis. lt appears that Ca v¢redzomus
not conspecific with C. dumetorum as believed by many earlier
workers, but rather a species which is ecologically allopat-
ric. Females' host acceptance, especially the preference for
Lotus scopartus by both species, indicates that the jtwoware
closely related... (It is possible that ¢€. virzdts iis a omeme
Specialized offshoot of C. dumetorum as suggested by the more
restricted distribution and acceptable host substrates mde
seen in the laboratory)... But as iC lench (1963) -assimeccmemEs
is also possible to think of C. virtdis as conspecific with
Callophrys shertdant (Carpenter) and because relatively little
has been recorded on the biology of the latter species, this
hypothesis remains to be tested. Further work of a biological
nature is needed to clarify confusing issues plaguing tnrs
subgenus.

Two species of California 29

TABLE 5 - CRITERIA USED TO SEPARATE THE IMMATURE STAGES OF
CALLOPHRYS DUMETORUM AND CALLOPHRYS VIRIDIS

STAGE CONDITION

C. dumetorum C. vtridts

MATURE LARVA:

a. Subdorsal white stripes present absent
b. Subdorsal ridges with absent present
red spots
e. Ground color green to yel- yellow-green
Ow green to whitish
d. Subdorsal and segmental absent present
ridges
PUPA
Color dark brown- light brown,
black mottled paler

Wing pad sculpturing pronounced weak

30

LITERATURE CITED

Barnes, Wm. and’ F. Hh.” Benjamin, 19235. "Callophrys dumetor-
um", Contrib. Nat. Hist. Lepid. North America, 5:64-69.

Billberg, C..J., 1820. Enumeratio insectorum in MaseomRaat-
berg, p. 0).

Boisduval, Jean, 1852. Lépidopterés de California. Ann.
SocsawEnt. France, ako) 229k

Brown, R. M. and P. A. Opler, 1967. Biological observations
on Callophrys viridis (Lycaenidae). J. Lepid. Soc.,
242 = ea,

Clench, H., 1944. Notes on lycaenid butterflies: a. The genus
Callophrys in North America. Bull. Mus. Conip. Zoeia
94(6):217-299.

1961. In Ehrlich &€ Ehrlich, How to nome
butterfites. Wm. €. Brown Co., Dubuque, pp. Z200= ae

1963. Callophrys (Lycaenidae) from the Pacific
Northwest... Jiu Res... Lep tidiis 9202) = 5 l- 160.

1966. Behavioral thermoregulation in butterflies.
Ecology, 47(6):1022-1034.

Comstock, J. A., 1927. Butterfiltes of Caltforntas) )heeauee
Printing Co., Los Angeles. Plate 50, pp 16o——oen

Coolidge, K. R., 1927. Life history studies of some Gatiger-—
nia Rhopalocera (Lepidoptera). '‘''The life history of
Callophrys dumetorum Boisduval''. Trans. Amer. Ent.

SOc...) 50(4) 3295 33)5.

Edwards, W. H.,1862. Descriptions of certain spectesmen
diurnal lepidoptera found within the limits of the Uni-
ted States and of British America. Proc. Acad. Wage
Sits ¢Phityla.g gh Wee oer

Ehrlich, P. R. and A. H. Ehrlteh, 1961... How to piers
butterflies. Wm. C. Brown Co., Dubuque.

Ford, E. B., 1945. Butterflies. N.N., Collins, Cone
ef Minny | 7215) NAS Il

Fracker, S. B., 1915. The classification of lepidopterous
larvae... (11. Biol. Monographs. 21): i201 22.

Gorelick, Gs A., 1970.2 A new subspecies of Callophrys
(Callophrys) dumetorum from Washington and Oregon (Ly-
caenidae). J. Res. Lepid., 7(2):99-104 ["1968"].

Henne, C., 1940. . Two new ispecies ot Lepidoptera from Gavan
fornia. Bull. So. Calif. Aead. iS mesa) 27 pee

31

Inter-Society Color Council, National Bureau of Standards,
Color-name chart illustrated with centroid colors,
sample #2106. Circular 553 (supplement).

Lawrence, D. A. and J. C. Downey, 1967. Morphology of the
immature stages of Everes comyntas Godart (Lycaenidae).
meres Lepid., 5(2):64 (["1966"]

McCorkle, D. V. (contributor), 1965. In: News of the Lepi-
dopterists' Society, ‘Annual summary''. No. 3, p 5.

Bene. , 1963. A California Flora. U. California Press

p- 849.

Newcomer, E. J., 1964. Butterflies of Yakima County, Wash-
magton. J. Lepid Soc. ,. 18(4):225

1965. In: News of the Lepidopterists'
Society (contributor), "Annual summary''. No 3, p 4.

mcrae 0. ,' 1968. Hilltopping. J. Res. Lepid., 6(2) :69-178.

Thompson, W. R., 1944. A catalogue of the parasites and

predators of insect pests. Imperial Parasite Service,
Bellevilte;- Ontario, Canada. Sec. 1, Pt. 5 (Lepidop-
tera) .

Tilden, J. W., 1963. An analysis of the North American spe-
cies of the genus Callophrys. J. Res. Lepid., 1(4):
281-300.

Wibtrams, FF. X.,

L910. Butterflies of San Francisco, Calhit.
Ent. News, 21(1)

36587 -

Ziegler, J. B., 1960. Preliminary redefinition of North
American hairstreak genera. J. Lepid Soc., 14(1) 19-23.

3) 2

APPENDIX
MATERIALS STUDIED

Callophrys viridts

Arranged alphabetically, 106 males and 80 females as
follows:

CALIFORNIA. Marin Co.: Point Reyes, 2M, 3F, IV-8-51
(C.D. MacNeill), 2M, IfI-30-56 (W.R. Bauer), 2F, V-16-58
(J. Powell:); Point Reyes, 9 mi. NE lighthouse, 4M) aaa
18-66 (R.S. Wielgus), 3F, IV-30-67, 2M, 8F, V-7-cny ef mee,
V-14-67 (all G.A. Gorelick). Mendocino Co.: Fort Bragg,
18.6 mi. N, 2M, VI-12-67 (G.A. Gorelick); Juan Creek, betw.
Rockport. and Westport on Hwy.1M, 4F, VI-9-67 (G.A. Gorelick) ;
Point Arena, 2M, 1F, VI-12-67 (G.A. Gorelick). San Francisco
Co.: Lone Mountain, San Francisco, 2M, 5F, III-6-17-1910
(F.X. Williams); Presidio, San Francisco 1M, 1F, IV-5-26 (no
collector), 1M, 1F, IV-18-29 (no collector), 1M, IV-10-38
(L.I. Hewes); San Francisco, 1M, V-28-05 (F.X. Williams), 1M,
IV-22-16 (K. Coolidge), 2M, IV=26-16 (K. Coolidge).
22-31. (A. Kusehe), 2M, JF, J¥-19-34, 3M, 1, Iv=27—9uee
IF, V-8-34, 1M, 2F, V-13-34 (all W. Hovanitz); Twin Peaks,
San Francisco, 1F, IV-4-60 (N. LaDue), 5M, 3F, IV-2-60 (R.
Stanford), 2M, 1F, LV-13-61 (P.A. Opler). San Mateo Co.:
San Bruno Mountains, 4M, 11-28-61, 4M, I1i1-10=60, 2usage-
III-17-61 (J. Powell), 2M, IITI-28-61 (N. LaDue), 1F, V-5-61
(J. Powell), 1F, III-9-62 (J. Powell), 1M, 3F); it —gaeee
(J.W. Tilden), 9M, 8F, Iv-6-7-62 (J..Powell) 3M, IV-10-62
(J. Chemsak, J. Powell), 2M, 2F, I1-26-28-63 (J. Powell),
IM, IV-28-63 (R.L. Langston), 2F, VI-4-63, emerged II-24-64,
TII-16-64 (reared from F. lattfolium latifolium) (J. Powell,
#63F8), 1F, ITI-17-66 (A.J“ Slater), 4M, 1121-19-66 (eene
Opler), 9M, 1F, III-25-66 (P.A. Opler, 1F, IV-13-66 (J. Pow-
ell), 5M, 2F, IV-16-66 (P.A. Opler), 4M, 1F, 1V—24=6e.mgeee
Wielpus), 2M, IlIl-2-67 (G.A. Gorelick), 2M, 11i—-7-og ee
TIT-22-67 (G.A. Gorelick).

Variant populations: El Dorado Co.: China Flat Camp-
grounds, 5 mi. S of Kyburz, 1F,VI-5-61 (N. LaDue). Seams
laus Co.: Del Puerto Canyon, 22 mi. W of Patterson, 2a
5-63 (R.L. Langston, J. Powell), 1M, 1h; 20-22-64. (eae
Langston, J. Powell, 1M, 1F, IV-6-49, in copulo (V.M. Stern)
(Adobe Creek); 1F, III-24-67 (G.A. Gorelick), 1M, IV-9-67
CP AsO plier):

Callophrys dumetorum

Arranged alphabetically, 270 males and 242 females as
follows:

BAJA CALIFORNIA. Baja California Norte: Tajo Branch of
Cantillas Canyon, Sierra Juarez, 1M, III-23-67 (J. Powell).

CALIFORNIA. Alameda Co.: Berkeley Hills, 1400', NE Oak-

33

land, 1M, IV-17-62, 1M, IV-15-64 (J. Powell); Oakland, NE,
IM, TII-9-30 (G. Heid), 1M, V-1-37, 2F, IV-9-38 (C.W. Ander-
son). Contra Costa Co.: Antioch, 4F, III-18-57 (P.A. Opler);
Antioch, 1 mi. E. 1M, III-19-60 (R. Stanford); Antioch, 1.5
mi, E, (Little Corral) 1M, IV-19-60 (P.A. Opler), 6M, 2F,
IV-20-67 (G.A. Gorelick; Mt. Diablo, 1M, IV-9-61 (P.A. Opler).
Fresno Co.: Coalinga, 23 mi. W, 1F, V-6-63 (R.L. Langston);
Huntington Lake, 1M, V-22-36 (L. Martin). Kern Co.: Havilah,
3000°', 2 mi. N, 1M, IV-28-64 (J. Powell); Walker Pass, 1 mi.
W, 1M, IV-26-64 (J. Doyen); Weldon, 16 mi. S, 7M, IV-26-64
(R.L. Langston, J. Powell). Lake Co.: Bear Creek, 3 mi. E,
en, Vi-24-67 (P.A. Opler); Middletown, 2500', 4 mi. WNW, 1F
V-15-58 (0.E. Sette); Whispering Pines, 3 mi. SE, 1M, IV-4-
62 (R.L. Langston). Los Angeles Co.: Altadena, 1M, III-7-26
(jee Comstock); Azusa, 2M, IV-3-45 (C.1I. Smith); Bob's Gab,
Gre Littlerock, 2M, IV-2-54, 1F, IV-7-55, 2M, 2F, IV-9-60
(P.A. Opler), 2M, IV-2-66 (R. Stanford); Cobal Creek Canyon,
eM, IITI-19-60 (K. Hughes); Desert Springs, 1M, V-5-56 (J.
Povey. Vi-/—-63 CR.L. Langston): Fish Canyon, 2M, III-7-=
B@eedomecotlector); Glendale, 3M, 1#, I11-26-44 (D.L. Bauer);
Griffith Park, 1M, IV-10-21 (J.A. Comstock), 2M, IV-14-60
(G.A. Gorelick); Mint Canyon, 1M, IV-16-23 (J.A. Comstock);
Pasadena, (b200' , 2F, LV-15-11°(F. Grinnell, Jr.): San Gabriel
Comome iM. TLI-1-25 (J.A. Comstock), 2M, III-17-61 (K.
Hughes), 2M, 3F, II11-20-67 (G.A. Gorelick); San Rafael Hills,
1M, IL-13-41, 2M, ITI-9-41 (D.L. Bauer). Marin Co.: Alpine
Lake, 1M, IV-25-58, 1M, IV-17-61 (J. Powell); Carson Ridge,
IM> 2V-16—-61. (J. Powell); Lake Lagunitas, 1F, V-2-59 (C.A.
iosens) Mariposa Co.: Bear Valley, 2.mi. N on Hwy 49, 1M,
ei 5—6i (PA. Opler). Mendocino Co.: Fort Bragg, 27 mv.
E, nr. Willits, 5M, 1F, VI-9-67 (G.A. Gorelick). Monterey
-Co.: Hastings Reservation, nr. Jamesburg, 3M, 1F, V-3-58 (J.
Powell); Partington Canyon, 1M, 1F, IV-21-51 (P.A. Opler);
Stone Canyon, 1M, 1F, IV-21-19 (E.P. VanDuzee). Napa Co.
Magia. NM, 2h, IV-19-45 (D.L. Bauer). ° Plumas Co.: Dixie
Game Refuge, 5800', 2M, V-17-64 (D.L. Bauer). Riverside Co.:
Hemet Lake, 1M, 1F, Iv-28-49 (C.D. MacNeill); La Sierra
College (campus), 1F, III-12-39 (D.L. Bauer); Lake Mathews,
south shore, 2M, III-19-67 (J. Lane); Nightingale, 1M, IV-28-
49 (C.D. MacNeill); Perris, 4M, IITI-4-53 (T.M. Blackman) ;
Sage, 5 mi. S, 2M, IV-16-65 (J. Pewell, D. Veirs). Sacramento
Cos: Brannan Island State Park, 50M, 150F, IV-7-to V-/-67
(G28. Gorelick). San Benito Co.: Bitterwater, 5.mi. S, 1M
mm-s0-59°(C.W. O'Brien); Hollister, 24 mi, SE, 1M, 3F, IV-
7-62 (0.E. Sette); Paicines, 5 mi. SW (Lime Kiln ‘Ra. ), 7M,
Pre et —24.-66 (A.J. Slater, J. Powell, R.L. Langston); Pin-
mnacles. Nat'l..Monument, 7 mi. W of Jct., 1M, IV-9-66 (P.A.
Opler). San Bernardino Co.: Desert Spring, 1 mi. E, 1M, 3F,
imgewi-o>5 (R.L. Langston); Fontana, .8) mi, n (Lytle Creek), 1F,
imemii=o5 (CA. Toschi); Highland, NE (City Creek), 1F, IV-17-
65 (G. Buckingham); Lucerne Valley, 5 mi. SW, 4M, 1F, IV-14-
64, 3M, IV-15-65 (R.L. Langston); Redlands, 1M, III-15-31
(C.M. Dammers); Upland, 1M, IV-9-57, 1F, IV-28-57 (T.M. Black-
man). San Diego Co.: Alpine, 4M, III-31-61 (R.L. Langston);
Alpine, 5 mi. E, IM, 1F, III-31-61 (J. Powell); Anza-Borrego
Desert State Park, 1000 Palms Canyon, 1M, III-4-34 (J. Creel-

34

man); Banner, 3 mi. H (Two Mile Hill), 2M, 1, I2i-23-3iam,
TII=22-42, 1M, 1F; TV-8-44,. 1M, TIIT-13-57, 1M, Lift-13=59) 20,
III-11-61 (F. Thorne); Banner, 7 mi. SE (Box Canyomn ems
TII-17-53 (Ff. Thorne); 1F, III-23-53 (3. Powell), Des@ameo
Ranger Station; IM, IITI-31-61 (R.L. Langston); EY Cajon
TIT-16-35 (CJM Brown); EL Cajon, 2.5 ami. Si 600 "aaa
TII-29-52, 1M, III-7-53, 2M, III-15-53 (F. Thorne); Jacumba,
3 mi. W, 1M,) T27-25-60 (R. Stanford), 2M, 1TL=2ieo ame
Thorne), 3M, ITI-22-66 (0. Shields); Lakeside, 2 mi. NE, a2,
TII-29-61, 2M, 1F, III-13-63 (J. Powell); <a Posta Creek
(Hwy. 80), 2M, IV-13-63 (PF. Thorne); Mission Gorgeous
TI-27-52, 2M, IIT=6=52, 3M, S2T21=52, 3M) 11292579
12-53, 2M, IT=14-53 (3. Powell); 1F, 1117-23-60 (Ry Seameaman:
Mount Laguna Jet.; 1 mi. N, 4F, “211-26-61 (R.L. vameeten.
BR.E. Lindquist); Mount Laguna Jet., 3 mi. N (Seeve "Gaara
2M, III-27-61 (R.L. Langston); Point Loma, 1M, Ii=taseame
II-12-35, 1M, II-17-35 (F. Thorne); Ramona, 9.5 mi. NNE
(Black Mountain, 4000'), 1M, V-9-65 (0. Shields); Sam Diego,
IF, IITI-4-33, 1M, I1T=16-33) VF) @i-12=35 (fF. Thomae ke
Warner's Hot Springs, 1M, IV-5-50, 2M, III-23-51 (J. Powell).
San Vicente Reservoir, 1M, III-19-49 (J. Powell); Scissor's
Crossing, 1 mi. W, 1M, III-28-53 (J. Powell), 2M, IV-3-60

(K. Hughes). San Luis Obispo Co.: Atascadero, 4 mi. anya
V-2-62 (J. Powell); Pozo, 6 mi. NE (Black Mountain, 3300'-=
3600"), 1M, LF, V=l~62 (CA. Tosehi; R.L. Langstom), gna

12 mi. NE (La Panza Camp), 1M, ,2F, IV=29-62 .(R.L. taneeeere
J. Powell); Simmler, 1M, III-20-40 (J.W. Tilden, G.S. Mans-
field). Santa Barbara Co.: Jalama Beach, 3M, 7F, IV-22-66
(R.L. Langston,’ J.) Powells Ava! Stater) >). Stas Cra. tee
Prisoner's Harbor, 2F, IV-29-66 (R.L: Langston; Sta. Ciaameu,
Upper Central Valley, 6F, IV-26-66 (R.L. Langston, J. Pow-
ell). Santa Clara Co.: Alum Rock Park, 1M, T1l-12-65 Ga.
Langston), 2M, IV-21-66 (P.A. Opler), 1M, TI-30-67 (GA.
Gorelick); Capitancillos Lake, 1M, T1T1-14-64 (PLA, Opleee
New Almaden, 1M, IV-17-64 (P.A. Opler); Palo Alto, 2M,

30° (W. Hovanitz): San Jose, 3M, IF; V-6-17 (K. Coole
Santa Cruz Co.: Boulder Creek, 6 mi. E, 5F, V-17-64 (PUA:
Opler); Mt. Ben Lomond, 2M, V-16-65 (P.A. Opler); Santa Cruz
Mtns., 2M, V-15-33') 2M, TTtu136) ‘4v=5=36 (5). idee
1M, V-1901 (J.G. Grundel), 1M, IV-—6-31 (no collector).

Sierra Co.: Sattley, 1 mi. W, 1F, VL-17-67 (G.A. Goreiaeee
Solano Co.: Green Valley, 1M, IV—8-53 (A. Telford). Goatees
Co.: Fairview, 9 mi. S,°3M, 1FP) IV-2/7-64 (W. Turners oo aeer
ell, P. Rude); Johnsondale, 2 mi.’ HE, IM, IV-27-64 (J 2) Poneeee
Sequoia National’ Park, 2M,-1F, 1V¥-16-30, 1M; V-16-30ume
Hewes). Ventura Co.: Gorman, 5 ‘mi. "S) Hunery Valley. eens
IV-10-60 (G.1. Stage): New Cuyama, 17, IIT228-57 |(R.P. idem).
sespe Canyon, 1F, IV-13-37 (G. Heid). Yolo Co.: Rumsey yuan
NW, 9M, 5F, IV-12-62 (2. Chemsakwy us. vPowel lye

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36

Fang.

Fags

FAirqh

EXPLANATION OF FIGURES

Eggs of Callophrys dumetorum on Lotus scopartus
bud, Brannan Is. St. Pk., Sacramento Gee
(G.A. Gorelick)

C. dumetorum larvae and pupa (same data as above)

C. dumetorum pupae (ventral, dorsal views), Pope
Creek, Napa Co. (J. Emmel)

Egg of Callophrys virtdis on Ertogonum lattifoltum
Lattfoltum leaf underside, Pt. Reyes, Marin Co.
(G.A. Gorelick)

C. vtridis larvae feeding on Lotus scopartius

C. vtridis pupae (ventral, dorsal views), San
Bruno Mountains, San Mateo Co. (J. Emmel)

+f

x

ed

38

Rigi.

14

EXPLANATION OF FIGURES

Callophrys viridis, dorsal view, Marin Co.
(G.A. Gorelick)

C. dumetorum, dorsal view, Sacramento Co.
(G. A. Gorelick)

C. dumetorum, same data as Fig. 8

C. dumetorum, ventral view, Sacramento Co.
(G.A. Gorelick)

C. dumetorum, same data as Fig. 10

C. dumetorum, ventral view, San Diego Co.
(J. Powell)

C, viridis, ventral view, Marin Co. (G.A.
Gorelick)

C.. virtdie: same.idata as Figi 13

39

40

Ears

Faingr

1
16

EXPLANATION OF FIGURES

Callophrys dumetorum egg

Mature, 4th instar C. dumetorum larva with
schematic dorsal view of abdominal segments
Vie vatial

Head capsule of mature C. dumetorum larva

Lateral view of 4th abdominal proleg showing
crotchet arrangement

Medial view of 4th abdominal proleg

Mature. 4th instar Ce viridis larva wien
schematic dorsal view of abdominal segments
Wee ete

C. dumetorum pupa, dorsal view

C. vtrtdis wing pad sculpturing (ventral view)

C. dumetorum wing pad sculpturing (ventral
view)

hi

«-
oo

ara
at 34)

My

d

o

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Comstock, J. A. 1927. Butterflies of California. Los Angeles, Calif. 334 pp.
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A SYNONYMIC LIST OF THE NEARCTIC RHOPALOCERA

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JOURNAL OF THE LEPIDOPTERISTS” SOCTER

The SUPPLEMENT series is intended for papers of
intermediate length, those which are longer than is normally
acceptable in the JOURNAL, yet shorter or of narrower
interest span than a monographic treatment which might
comprise a MEMOIR. The SUPPLEMENTS are issued at irregular
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This issue edited by J. A. Powell.
Supplements to Volume 23 (mailed May 6, 1970):
1. A review of the Megathymidae; by H. A. Freeman; 59 pp.

2. Records, new species, and a new genus of Hesperiidae
from Mexico; by H. A. Freeman; 62 pp.

3. On the frequency of butterflies in eastern Brazil, with
a list of the butterfly fauna of Pocos de Caldas, Minas
Gerais; by Hans Ebert; 48 pp.

Volume 25

JOURNAL

of the

" Lepworrenisrs SOCIETY

"Published Akay by THE LEPIDOPTERISTS’ SOCIETY

" Publié par LA-SOCIETE DES LEPIDOPTERISTES |
_ Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN

VOLUME 25

SUPPLEMENT 3

BIOLOGICAL STUDIES ON MOTHS OF THE GENUS
ETHMIA IN CALIFORNIA
‘(Gelechioidea)

a

by Jerry A. Powell

10 May 197)

THE LEPIDOPTERISTS’ SOCIETY
Se all va) EDITORIAL COMMITTEE. 3

Mame ey oe Hardwick, Editor of ane Tourkal’. AN Oy ice
CV's icavelt, Editor lof. the (Weve; =! Cancer
rs ‘oft 1A Hesse) ; Manager) of the Memotrs | Py rte aa
By RIE Fay EXECUTIVE CoUNCIL GOAL SN ORO | en
Ne se ke Pan (New Haven, Copy y; Prestaeat PS Hey?
sb. M, Martin (Prescott, Ariz.), President- elect» |
dae Freeman’ (Garland, Tees) 4 Ilst\Vice- President
wlian Jumalon (Cebu City, Bip phneet 2 Vice- ‘President
.W.. Philip (Fairbanks, Alaska) , Vicer President aa
S$. Nicolay, (Virgihia Beach, Va.), ‘Treasurer
ee (CédariFalls, We. ), Secretary” & ROA.
: ‘Millen (Sarasota, Fla. as Secretary - ~eleeh i) 9 so ales

Wenbers ae Lorde (three, year term) M. Ogata 1972
Sr Brower: 19OPD wii.’ -E.C. Welling el

Z pa “McGuffin: 1971 i - Andre. Blanchard 273
| Nekrutenko, TOV ie {ORB bomtnick. 1978 ‘
ee Mather OAR A ‘ AEA PO ME ae Sey oy

i % " * 5 i lide * * o-* ; A

The object of The Lepidopterists' Sopiety, Whee was _
formed in May), 1947, and formally constituted in December 1950,
is "to promote: the ‘science of lepidopterology in all its |
‘branches, ja ee. cis Suave periodical and other publications ~
on Lepidoptera, to facilitate the exchange of. specimens and
ideas by both the professional worker and’ the amateur in the
field; to secure cooperation in all measures"! HERS aby aa

towards these aims. ye i

‘Membership. “Ty” the Society is open to aN persuus kntenee esa a
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‘ November, and eight. pom > of Ene News each year. hy, ,08 ae

=r ‘ ; ae S SE.
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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY

VoLume 25

SUPPLEMENT 3

BIOLOGICAL STUDIES ON MOTHS OF THE GENUS
| ETHMIA IN CALIFORNIA

(Gelechioidea)

Jerry A. Powell

University of California, Berkeley

TABLE OF CONTENTS

Introduction

Techniques

.Acknowledgements

Ethmta coqutllettella (Busck)

Ei

if -

E.

Ee

scylla Powell

brevtstriga brevistrtga Clarke
b. ardteola Powell

albttogata Walsingham
plagtobothrae Powell

minuta Powell

charybdts Powell
albistrigella (Walsingham)
nadta Clarke

semtlugens (Zeller)
arctostaphylella (Walsingham)
discostrigella (Chambers)
semttenebrella Dyar

timberlaket Powell

Literature “Cited

Index to host plants

l[llustrations

INTRODUCTION

The family Ethmiidae is composed of small to moderate
sized moths and is world-wide in distribution, with its
greatest diversity occurring in the Neotropical Region.
Consisting primarily of the one large genus Ethmia the
group is distinct in many respects, without close relation-
ship te other families. Ethmiids have in the past been
considered as related to or members of the Oecophoridae.
Probably they are most closely related to the Stenomidae,
and the three groups are considered to be families in the
Gelechioidea by present workers.

About 30 members of the genus Ethmta have been reared
previously, primarily in the Palearctic and in the eastern
United States. Nearly all feed externally on Boraginaceae
or Hydrophyllaceae during the larval stage, but there have
been few detailed studies. Habits of the few other Nearctic
genera are equally poorly known: species in Pyramntdobela
feed on Penstemon (Scrophulariaceae) and Buddleta (Logani-
aceae) (Braun, 1921; Keifer, 1936), while the biology of
Pseudethmia is unknown. Two other species formerly consid-
ered to be ethmiids, Bumeyricktia trimaculella (Fitch) and
"Ethmia" eoltoradella (Walsingham), are fungus feeders and
have recently been transferred to the Oecophoridae, a group
containing genera with similar morphological and biological
Easy tbawrence and Powell, 1969).

In connection with a California Insect Survey project
on Ethmiidae, | began to investigate the biologies of these
moths in 1961. The study gradually developed into a com-
prehensive taxonomic one encompassing the New World fauna,
some 135 species. Field efforts were particularly directed
‘towards Ethmta in California, resulting in the present data
on 14 species, which, however, represent only a few species
groups in one section of the genus. Thus it seems appropri-
ate to give this detailed biological information separately
from the systematic treatment of the genus as a whole. A
general review of biological knowledge for the family is
Biveneine that study (Powell, 1971).

lt became evident early in this work that some species

are diurnal and others nocturnal. lt was one of the aims

of the investigation to clearly define which are diurnal in
Sidem fo assess..the. significance of this phase of the moths
biology in systematic relationships. Adults of both diur-
nal and nocturnal species are sometimes encountered in num-
bers during the daytime, and at times it is difficult to

distinguish between active flight behavior and reactionary
movement in response to disturbance by the observer. There-

1 This study was in part made possible through assistance
from National Science Foundation Grants GB-4014 and
GB-6813x, "Comparative biology in relation to systematics
of Microlepidoptera", 1965-1970.

k Introduction

fore, criteria | have used for defining natural activity

rhythm in the diel cycle have been the timing of mating,

oviposition, and the "quiescent posture'' by moths in cap-
Livi by

The resting posture during periods of activity was
distinguishable from that shown by moths in the !''quiescent
posture'' assumed during the inactive phase of the diel
cycle. In the ''quiescent posture'' the wings were tightly
clasped against the abdomen and the antennae were held
back alongside the body, under the costal edges of the wings.
The insects crouched low, almost appressed to the substrate,
with the legs widely outstretched. Moths temporarily not
“moving during activity periods held the wings somewhat
loosely spread from the sides of the abdomen, and the an-
tennae projected outward, at right angles to the body axis
or somewhat forward, usually moving slowly. At the same
time they stood up higher above the substrate with the legs
less widely outstretched.

The ''quiescent posture'' was exhibited at night by all
individuals of species which mate and oviposit in daytime.
There was not a temperature correlation with darkness, since
heating indoors kept the temperature above 18° C until mid-
night or so, while it often remained as low as 12° during
the early daylight hours.

As a result of this study, it is now known that diurnal
species possess small eyes and usually very dark integument
and vestiture. Thus the behavior pattern can be predicted
on the basis of preserved specimens. The eye size has been
quantified and described elsewhere (Powell, 1971). In a
few species the eyes are intermediate in size, and my obser-
vations on Ethmta arctostaphylella suggest that this is
correlated with a tendency towards crepuscular behavior.

TECHNIQUES

The present data originate from field collections of
either late instar larvae or adults which were retained alive
for oviposition. The moths were taken at lights or by net-
ting them during daytime or at dusk. Adults were transported
from the field in cotton-stoppered glass vials and were
caged in one-gallon glass breeding jars. The housing methods
and details of the container were essentially the same as
used and described in studies of tortricine moths (Powell,
1964). The jars proved more satisfactory for ethmiids than
for Tortricinae because oviposition by *#thmta usually occur-
red on the host plant or on the nylon mesh used as a ceiling
on the cage (figs: 11, 12), rather than on the sides onueqe
glass container. Insofar as possible the jars were placed
adjacent to an open window, exposed to natural lighting,
including direct, dappled sunlight (filtered through tree
foliage) during part of the day. Observations after night-
fall were made by means of a flashlight provided with a
cover of red construction paper. This light source usually
did not attract or otherwise disturb the moths.

Biological studies on Ethmia 5

In the field larvae were generally detected by hand-
searching suspect plants. A few species could be effective-
ly collected by beating (certain perennial plants) or sweep-
ing, but most of the present species live in concealed
shelters on low, herbaceous hosts. Larvae were transported
from the field in polyethylene bags and generally were
housed in closed containers with cut sprigs of host plant.
Newly hatched larvae in the laboratory were usually housed
in 25 x 100 mm salve tins on small bouquets of foodplant in
water vials plugged by cotton. In few cases greenhouse
plants were used for early instar establishment. Larger
larvae were housed in plastic sandwich boxes or 85 x 100 mm
jars with cuttings of foodplant.

To provide fresh plant material for species reared from
eggs, plants from the collection site were transplanted to
pots in a greenhouse, bouquets were placed in water, or cut-
tings of the same or a closely related plant were obtained
from the University of California, Berkeley, Botanical Garden
at the time of egg hatch. Transplanting (perennial Phacelia)
or cuttings in water (annuals) proved satisfactory for early
instars since these plants usually persisted well for 2-3
weeks. However they matured in advance of field conditions,
and later instars were provided with plants from the Botan-
ical Garden unless the original collection site could be
revisited conveniently. For field collected late instar
larvae, cuttings were either offered as bouquets in water or
were refrigerated (+ 4° C) and later offered without water.
This required frequent replenishing of provisions.

Soft paper toweling, folded many times, and sometimes
cut sections of dry Yucca whtpplet floral stalks were pro-
vided as pupation substrates. A tendency to wander and
burrow into soft bark and similar substances has been repor-
ted in the literature for several Ethmia.

Rearing was conducted at laboratory temperatures (usual -
ly varying about 12-20° daily). Pupae in diapause were in
some cases housed in an outdoor screen cage at Berkeley or
in an open shed at Russell Property, near Lafayette, Califor-
nia, an inland station where greater climatic extremes more
approximate field conditions of inland parts of California.

The only previous biological information concerning the
species discussed below is the report by Dyar (1902) that
E. semttenebrella had been reared from Cercocarpus in Ari-
zona. Hosts of three species must have been known to Keifer
(1936) who described the pupa of albitogata and mentioned
larval characters of two other species, but he gave no
information on their biologies. Therefore | attempted to
discover the host association through observing moths in the
field. Adults of both diurnal and nocturnal species tend to
stay close to the larval foodplants, and each of the present
species has proven to be specific to members of one plant
genus so far as known. Following field collection, |! caged
females with a small bouquet of the suspect host. lf no
clear association had been ascertained, a varied menu of

6 Introduction

possible oviposition substrates was offered. Females exhib-
ited only a poor oviposition response or none at all, if
caged without the appropriate host. The fact that most
previous recorded plants for Ethmia are Boraginaceae and
Hydrophyllaceae helped to restrict my selection, during
field searches. However, in one case this restricted think-
ing hindered the eventual discovery of an unrelated plant

as the host.

Detailed comparison of eggs was not attempted. Photo-
graphs showing general habitus and placement on the plant
were taken for most species. High magnification scanning
electron micrographs were executed only for ecy?7ae) eam
scheduling for photography eggs were usually stored in a
refrigerator (4° C) for several days, which delayed matura-
tion of developing larvae for a period about equal to that
in cold storage.

When sufficient numbers were available, larvae repre-
senting each instar were preserved, using KAAD for a few
minutes, followed by storage in 95% ethyl alcohol. Head
capsules representing previous instars of living individu-
als were recovered and were used in measurements for esti-
mating the number of instars, along with the preserved
larvae.

Detailed morphological descriptions of the larval
Stages have not been made. The larvae are briefly charac-
terized, with special reference to instar differences,
following the biological discussion of each species. Abbre-
viations are as follows (see fig. 7): HC = Head Capsule, a
measurement of maximum width as seen from above is given;
ThSh = Prothoracie Shield; Pin = Pinacula; D = Devout
a narrow color stripe in later instars of Most Specrean
from pro- or mesothorax to ninth abdominal segment; DL =
Dorsolateral pigment bands tateral to D, above the spieaetes:
L = Lateral band, a broad area around and below spiracles;
LV = Lateroventral band, a weakly developed pigment band
below L, above the legs; AbdCr = Abdominal Crotcehets seam
extended mesoseries or mesopenellipse in all the present
species; AnCr = Anal Crotchets. All measurements were
made through a disc micrometer at 27x or 54x magnification
and are given in mm, based on specimens distended in KAAD.
Measurements are based on six or more specimens except
where indicated otherwise, with the number in brackets [ ].
Color features were noted from living larvae. Integumental
colors of Ethmita fade during preservation.

Morphological characteristics of the pupae are varia-
ble, and only limited comparison between species is attemp-
ted owing to inadequte series for most species. Most of
the species examined here are quite similar.

Biological studies on Ethmia 7

CONSTANT BIOLOGICAL CHARACTERS

Several features of the biology and behavior appear to
be consistent among all species, and these are not discussed
for each species. The eggs are deposited singly, cemented
to the substrate by an affixed area nearly as long and wide
aonone egg. The egg is more or less rectangular-.in outline,
nearly as thick as wide; it does not flatten out onto the
substrate, but conforms to minor irregularities, sometimes
aeering its shape a little (e.g., fig. 12). At hatching
larvae chew a round, ragged hole at the micropylar end,
sometimes well off center. The hole is about one-half the
diameter of the egg, and no further feeding is done on the
eggshell by newly hatched larvae. When disturbed or dis-
lodged, larvae of most species, especially in later instars,
react by wriggling violently backward. A few species feign
death and fall, immobile, to the substrate. Based on head
capsule measurements, there appear to be five instars in
several species, but data are too fragmentary to determine
the number with certainty for most species. At least in
brevtstrtga and secylla, and probably plagtobothrae and albt-
togata, there are five, while large species such as discos-
trigella and arctostaphylella probably undergo six, at least
winewsione individuals (figs. 1-6). At maturity ltarvae of
probably all species wander, often having been found to bur-
row imto soft, woody substrates to pupate. Data on speci-
mens | have examined, especially of several species reared
Soe Parker—at the University of California, Davis, from
trap nests!, indicate this is a widespread habit. During
my study larvae usually were confined in salve tins or 35mm
pill boxes for pupation, and most did not burrow into yucca
Puen using unnatural situations such»as a corner of the
container. At emergence the pupal shell remains inside the
cocoon, held in place by the hooked setae of the anteriorly
directed “anal, legs! (figs..8, 9) (in all species except
seylla). The cremaster such as is normally developed in
most Lepidoptera at the tip of the abdomen is vestigial.
Eight frail setae are present in a constant arrangement for
all species, but these do not aid in anchoring the pupa.

The degree to which the "anal legs'' of the pupa are appressed
to the abdominal venter or angled outward varies within
species, possibly affected by the shape of the cocoon.

ACKNOWLEDGEMENTS

Grateful acknowledgement is made to Helen K. Sharsmith,
formerly of the University of California, Berkeley, Herbar-
ium, who provided many of the plant identifications associa-~

1 Sections of Sambucus stems, 45 cm in length, which had
been stuck into the ground, with a 1.5-4 mm hole drilled
in the exposed end (Parker and Bohart, 1966).

8 coqutllettetla

ted with this study. “Throwgh Mrs.) Sharsmith, Linealnepen—
Stance, University of California, Berkeley, determined most
Of the Phacelia, and Francia Chisaki Hommersand, Chapel Hill,
North Carolina, identified some of the Amsinckta species.
Thanks are due Anton Crist and other personnel of the Uni-
versity of California Botanical Garden, who have been coop-
erative in providing cuttings of native host plants many
times during the past decade. Photographs of eggs in sttu
were executed by A.A. Blaker, Scientific Photographic Labor-
atory, University of California, Berkeley. Stereoscan Scan-
ning Electron Microphotographs were executed through the
cooperation of T.E. Everhart, Electronics Research Labora=
tory, University of California, Berkeley, through sSuppore
from National Science Foundation Grant GB-6428 and Grant

No. GM15536 from the National Institutes of Health. Assist-
ance with field collections, particularly during early years
of the study, was given by several persons, whose help is
appreciated and acknowledged by mention in the text; but
special thanks are due C. Don MacNeill, Oakland Museum, and
Catharine Toschi Tauber, Ithaca, New York. Where not other-
wise indicated, all collections and observations are my own.

ETHMIA COQUILLETTELLA BUSCK

Ethmta ecoquittlettetla Buseck, W907, Proc.’ Ent: SOG w Wamu
8:95.

This species has been collected at only a few widely
Scattered sites in arid parts of California and interior
British Columbia during the 70 years since the original
specimens were taken in the vicinity of Los Angeles (Powell,
1959,°1971).. "Considerable interpopulational variatvon se
exhibited, and study of more material will be necessary in
order to confirm. that’ just one species Is Involved.) Same
and related species in the southwestern United States, which
have the palpi clothed with stiff, erect bristles, are’ be=
lieved to be diurnal.

| have not been able to confirm the foodplant of coqut-
tlettella with certainty. The moths have been encountered
only in small numbers, and not in close association with any
plant. Larvae accept Phacelta and Nemophila in the labora-
tory, and on two occasions have been reared to the fifth
instar but not to maturity. One larva was collected in the
file Wd on Priacel ta dvs tones

Study areas: - 1)Pinyon Flat, 16 road miles southwest of
Palm Desert, Riverside Co.; 1 male, 3 females taken in
fFlight,°11200 AGM: =<"? 700°PEMe, "Apri? 13571963" (Cl Al tocrem

and J. Powell), 3\ females retained alive (63D19) .” 2) anne
road Canyon, 4 miles northeast of Elsinore, Riverside Co.;

| male, 2 females at flowers of Coreopsis californica (Com-
positae)); ( 11:45°A.Ma.-92 00) PME Js hApmiWols, 196 Seige
Toschi and J. Powell), 2 females retained! ative (650m

3) Del Puerto Canyon, 23 road miles west of Patterson, Stan-

Biological studies on Ethmia 9

1slaus Co.; 4 males, | female taken in flight, 12:30 - 3:00
Pateanetcagen 25, 1969, 2 males, 1. female retained alive
(69094); 1 larva on Phacelta distans, April 27, 1969 (69D59).
At Pinyon Flat the moths were taken near Mentzelia (Loas-
aceae), a possible nectar source, and immature Phacelia
ditstans var. australis; at Railroad Canyon a mixed stand of
Phacelta eteutaria var. hispida, P. distans, and Nemophila
menatestt grew near the collection site; while at Del Puerto
Canyon, Phacelta distans and Amsinckia tntermedta were sus-
pece uhos ts.

Adult behavior: - Collection records indicate a single
Piughe period in early spring, in California from mid Feb=
ruary to mid April, varying with conditions of locality and
Season. Laboratory observations have been sporadic but tend
to confirm the diurnal behavior pattern indicated by adult
morphology and field collections. In the breeding jar moths
were active during the day and occasionally at night if
direct lighting was on them. Mating was not observed.

Females from Pinyon Flat were caged with Phgceliq and
Mentzelita tn a 85 x 100 mm jar, but on the day following
collection the jar became water soaked during transport in
a field ice box. Two of the females recovered and were
transferred to a one-gallon jar; but probably they had been
weakened as it appeared that neither left the floor of the
container. Only 9 eggs were deposited, on the cardboard jar
Floor.

Females from Railroad Canyon were aiso housed in an 85 x
100 mm jar under field conditions. A bouquet of Phacelia
eteutarta, Nemophila, and a Coreopsis flower was offered as
possible oviposition substrate and nectar source. A natural
photoperiod rhythm was not provided, and, with exposure to
indirect and direct lighting late in the evenings, activity
periods apparently were irregular. A total of 40 eqgs was
deposited by one'or both females, in part during early hours
OF mugnte (8:00 P.M. - 2:00,.A.M.) and during early morning
hetimsec2 00) A.M.. - 8:00 A.M.) . .A few eggs were place on
Leaves of Nemophtla but most were deposited around corners
of the container floor.

The adults from Del Puerto Canyon were caged in more
suitable conditions, in a one-gallon breeding jar provided
with a bouquet of immature Phacelta dtstans, Amstnekta in
flower, and immature Eptlobtum sp. (Onagraceae), housed in
natural photoperiod. Males lived only 1-3 days, but the
females survided 10 days, depositing 70 eggs, nearly all
during the first two days of confinement. Both sexes were
inactive at night, remaining in quiescent posture until
10:00 - 11:00 A.M. The period of highest activity appeared
to be 1:00 - 3:00 P.M., although individual movement occur-
red in dappled sunlight until 6:00 P.M. The moths displayed
a stronger positive phototropic response than some other
diurnal species. Oviposition was observed between 1:30 and
3:00 P.M., and a few eggs were deposited later in the day.
The female selected roughened surfaces in the side of the

10 coqutllettella

jar towards the light. She walked about on undersides of
Phacelta leaves and on the nylon screen ceiling with the
abdomen extended and curled downward, probing at the sub-
strate. Usually she ran a few ''steps'', then probed two or
three times, sometimes slightly to the side. A quiescent
pause of several seconds followed each egg deposition.

Of the 70 eggs, about 75% were deposited around the s
side of the jar towards the light; and of the total, 60%
were placed on the masking tape around the floor and 15% on
the nylon. Only 8 eggs were deposited on the plants, all
on the undersides of Phacelta leaves adjacent to the light
side.

Egg. - Eggs were uniformly subrectangular, varying from
0.46 x 0.76 to 0.41 x 0.82 mm in outline. During develop-
ment all turned pink by the third day, and a somewhat dark-
er reddish by the 7th to 9th day. Incubation time varied
from 10 days in April°in the fleld (6501) to’ 11-12 edawemn
March at room temperatures (69C94). Emergence sometimes
occurred well off center of the micropylar end.

Larva, - Considerable difficulty was encountered in
inducing young larvae to establish and feed, relative to my
experience with other Ethmia. A distinct preference for
flower buds was shown, and it appeared that leaves were un-
suitable for development. A continuous supply of immature
flowers was not provided, and this may have been a critical
factor in the failure of larvae to reach maturity under lab-
oratory conditions.

First instar larvae (63D10) placed on Phacelia distans
from Pinyon Flat which had been,in refrigeration 16 days
failed to survive. Some fed.a little at the base of buds,
but none successfully established themselves.

One and two day old larvae (65D1) were placed on fresh
buds of Nemophitla maculata and Phacelta tanacettfolta from
the Botanical Garden; larvae at first began feeding on both
plants, either in the buds or in crotches of leaflets or
sepals. All eight larvae placed on the immature, scorpioid
spikes of the Phacelta established successfully and reached
at least the second instar. The inflorescences were tightly
curled, and by the second day frass was visible between the
appressed, hirsute buds. Larvae continued to feed inside
the buds during the first 12 days; on May 7,10 day old waiam:
material from the Botanical Garden was added when the larvae
were 11-12 days old. Three days later only three larvae had
moved to the more recently offered buds. On May 14, the 3rd
and 4th instar larvae were moved to fresh Phacelia tanacett-
folta from the Botanical Garden. By this time the plants
had bloomed and subsequent feeding took place mainly on
smaller leaflets, often those adjoining flowers.

The ephemeral character of the Wemophila flowers preven-
ted establishment of all but one of the first instar larvae.

Biological studies on Ethmia 11

This larva succeeded in burrowing into an unopened bud, pre-
venting it from further development, and fed on the pistil
and stamens for 11 days; it then migrated to a new, less de-
veloped bud and began feeding. The 15 day, 3rd instar larva
left the Nemophila (which was partially collapsing in the
Heer wial)-and was transferred to the Phacelia. At the
Botanical Garden the Nemophitla was drying by the time the
larvae were 19-20 days old, which, together with the diffi-
culties encountered in establishing on this plant, suggests
that Nemophila is not a suitable host. Nutritionally the
Nemophtila flowers appeared to be adequate since the one lar-
va waS as mature as the most advanced of those feeding on
Phacelia at each inspection.

On May 24, about 30 days after hatching, laboratory ob-
servations were interrupted by a vacation camping trip.
The remaining larvae were carried in a salve tin and were
subjected to greater temperature fluctuations than in the
laboratory and to drying of the foodplant. Fresh Phacelia
leaves were added from San Bernardino County, California,
and Coconino County, Arizona, but the final larva died by
June 4 while an early fifth instar.

Unfed first instar larvae (69C94) were placed on Pha-
celia distans from Del Puerto Canyon, which had been kept
in water 12-13 days; establishment was affected by burrow-
ing into unopened buds. After 4 days an accumulation of
fine frass was noted in the dense hairs of the inflorescence.
On Aprit 19, the 10-12 day old 2nd and 3rd instav larvae
were transferred within their shelters to vials with fresh
Nemophila menztesti from Santa Clara County. Although the
original Phacelita had become blackened and mouldy, after 3
days there were no signs of feeding on either flowers or
Weawes! of the Wemophtia. The 13-15 day old; 3rd instar lar-
vae were then offered fresh Phacelta tanacetifolia from the
Botanical Garden, and all established new shelters. Subse-
quent feeding occurred in buds on bouquets of this host.
Fresh sprigs were added every few days, as it did not keep
well in water, and two exposures to badly withered plants
probably affected larval development. Owing to a field trip
intervention, surveillance was terminated on May 9, and the
30-32 day old larvae were preserved. They apparently were
penultimate and immature last instar individuals which had
not grown during the preceding 4 days due to the condition
of the plant.

Inspection of Phacelia distans at Del Puerto Canyon on
April 27 (when laboratory larvae were about three weeks old)
revealed only one larva of coquillettella. This individual
was provided P. ditstans for 8 days, then P. tanacetifolta.
However, foodplant conditions were intermittently poor, and
the Warva died on May 7°in*the final instar.

Since feeding took place on buds of Phacelia tanaceti-
folia in water and the developmental rate was similar to

12 seylla

that of Ethmia brevistriga, it is assumed that P, distansand
P. eteutarita might serve as suitable hosts at the study sites.
However, as with other species in the diurnal group, a defi-
nite preference for feeding in unopened flowers was shown,
and it may be that fresh buds are necessary to provide suffi-
cient nourishment to complete development.

First instar: Length 1.6 mm; HC 0.20-0.22 mm, brown, ocel-

lar area black; ThSh brownish; integument and setae unpig-
menved.

Seeond instar: Length [1] 4.0 mm; HC [4] 0.30-0.35 mn,
brown, frontal area slightly paler; Thsh, thordei¢geieze
lateral spots on prolegs, and anal shield, browns Pamieeeuce-
Say integument otherwise unpigmented. AbdCr 8-10; AnCr
La ad we pers ¥

Third instar:none preserved; HC and ThSh dark brown, in-
tegument pattern pale purpiasne:

Fourth instar: Length [2] 8.2-8.5 mm; HC [3] 0.67-0.74
mm, brown, mottled with pale areas; ThSh mottled, brown with
darker spots; Pin blackish; D white (not unpigmented) with
thin, median deep ochreous streaks; DL mottled, pale purplish
to brownish olive, large, distinct white rings encircling

pinacula; AbdCr 7-10 (usually 8-9); AnCr 7-8.

Fifth instar: [5] Length 8.0-12.0 mm (none mature);
0.74-0.82 mm, markings strongly contrasted; integument oat
ors similar to penultimate, ThSh brown with darker spots;

D and pinacula rings of DL more contrastingly white; See

darker purplish or olive, ale sliche ly gwhicssh jex unpigmented,
With or without #aing ochreous streak; AbdCr 9-14 (usually
12S) We NaC eile

ETHMIA SCYLLA POWELL
Ethmia seylla~ Powell, 197, U. Calif. Publ. Bnt of) se pees

Three localities along the inner Coast Range of central
California comprise the known range of seylla. 1 collected
the first specimen on March 18, 1959, at about the time my
review of the poor state of knowledge of Ethmta in Califor-
nia had gone to press (Powell, 1959). However, exactly ten
years were to elapse before |! was able to solve the mystery
of scylla's biology, the search having been hampered by a
preconceived notion that some Borage or Hydrophyll must be
the hostplant. Although the adults resemble £thmia brevt-
striga, and thus might be expected to feed on a Phacelia,
this species proves to be unique as the only member of the
genus known to use Scrophulariaceae and further the only
member of a Nearctic or Holarctic species group which does
not possess the peculiar ''anal legs'' of the pupa.

Study dreds’. = 1)Russelmann Park, north slope of Mt.

Biological studies on Ethmia 13

Diablo, 1100 feet, Contra Costa Co.; adults numerous, April
2, 1960 (a... Burns? and’ J&0 Powell): adults Sparsies: April 6;
1962, 3 males, 2 females retained alive (62D2). 2) Raines
Park, Del Puerto Canyon, Stanislaus Co.; adults sparse,
March 5, 1963,(R. Langston and J. Powell), 4 males, 2 fe-
males retained alive (63C1); negative results checking Am-
stnekia tntermedia, April 30, 1963; adults abundant, March
19, 1969, 4 males, 3 females retained alive (69C90); adults
common, March 25, 1969, 2 females retained alive (69C90);
larvae abundant in flowers Collinsta heterophylla April 27,
1969 (69D60). 3) Three miles northwest of Rumsey, Yolo Co.;
adults sparse, March 8, 1964.

Adult behavior. - The species has a single annual flight,
in moderately early spring, from late February to early
April, varying two to three weeks with seasonal conditions.
This is well ahead of the bloom period of the foodplant.

All. three localities are deciduous oak-digger pine scrub
forest situations. At Russelmann Park the moths appeared

to fly around poison oak, Rhus diversiloba, in the manner of
Ethmta albttogata at the San Bruno Mountains, where Am-
stncekta grows in close association with the poison oak
clumps. Adults of seyllqa sometimes perched on poison oak
foliage where they resemble bird droppings. The association
at Raines Park proved to be a general one; the Colltnsia is
abundant in semi-shaded spots on north slopes around various
shrub growth including Juntperug, Arctostaphylos, and
Ceanothus.

Adults were observed in the field between 11:00 A.M.
and 4:00 P.M. Two mating pairs were taken, one at 3:40 P.M.
oulAenin 2; 1960.° Mating did not occur in the laboratory.

In captivity moths intermittently abandoned the quies-
cent posture between 8:30 and 10:30 A.M., but continuous
Settumey did not begin before 11:00. It lasted until about
k:00 P.M., after which movement gradually subsided, ceasing
byegetser 6:00 P.M., at about sunset. As indicated by>ovi-
position, the height of activity was not concentrated into
a brief period and did not strongly vary between cloudy and
clear days, extending from 12:00 or 1:00 P.M. to about 4:00.
Generally adults were strongly positively phototropic, and
oviposition by the 1962 females seemed to corroborate ERTS...
However, when Collinsta was offered (69C90), eggs were more
evenly distributed, with more than half of those deposited
on plants having been placed on a Collinsia in the center of
Bie,jat, rather than on those nearer the light source.

In earlier collections various immature, low growing
annual plants from the collection sites were included in the
breeding jars. The 1962 lot was also provided with a few
small Plagtobothrys nothofulvus from Lake County. Several
small Amstnckia intermedia in bloom were included in the
1963 trial. One or more of the 62D2 females deposited 5
eggs on the nylon ceiling and 30 on the cardboard floor
(which was roughened, with fibres protruding, the result of

14 seylla

removal of masking tape), 80% concentrated on the side of
the jar towards the light. In experiments with other #eh-
mta the cardboard floor was sometimes used by old, weak fe-
males, but in this case it was selected on the first day of
confinement, judging from the incubation period.

The 1969 moths were offered an array of Amsinektia in
early bloom, and immature plants of both Phacelta distans
and Collinsta heterophylla. The latter had been observed
in high numerical density in the vicinity of female moth
concentration. In the breeding jar females complied by dis-
playing a distinct selection not only for Collinsia, but
for the leaf axils. A total of 124 eggs was deposited by 4
females; exactly half were placed on the nylon screen, while
nearly all the remainder (60) were laid in leaf axils of
Collinsta. Most were on the upperside of the petioles, but
some were placed in axils where secondary leaves originated,
so that they were sometimes affixed to undersides of secon-
dary petioles when tucked into the primary axils. They were
distributed along the height of the plant, but tended to be
concentrated (60%) on the middle axils, which bore the sec-
ondary leaf growth. One egg was deposited on the upperside
of a leaf blade. One was deposited on each of the Amsinckta
and Phacelta.

Females tended to concentrate on the screen ceiling
towards the light source, but wandered during oviposition.
The probing action of the extended ovipositor was more or
less continuous at about 30/minute. One female was observed
in this behavior on the nylon, traveling some 7 cm during
2.5 minutes; finally after 4 minutes she extruded an egg
onto the apparently uniform nylon mesh. Deposition of an
egg required 1-2 seconds, after which the female usually
quickly moved several cm without probing the ovipositor,
then stopped in a stationary pause, sometimes moving to the
light side to do so. Results of individual female's efforts
were not tabulated, but 3 females deposited a combined total
of about 100 eggs in 2 days.

The moths did not survive long under cage conditions,
males living 3-7 days, and the females only 3-5 days.

Eqg. - (figs. 13-17)) The: egg is characteristical lyneuom-
gate, cylindrical, with the chorion strongly reticulated
with ridges which bear no microstructural modifications.

Eggs varied! from 0.3:00* 0.59 mm t0.027 ix 056 1imme

During development the eggs changed color, to pale or-
ange by the 2nd day, bright pink by the 3nd) day, andigpaqu-
ally darker reddish before the larvae became visible prior
to emergence. Incubation required 10 days (62D2) and 8-9
days (69C90) at laboratory temperatures.

Larva. - In order to confirm the host selection dis-
played in oviposition behavior, separate lots of newly
hatched larvae (69C90) were segregated in 32 x 90 mm shell
vials and were offered cut terminals including flower buds,

Biological studies on Ethmta 15

of four menus: a) Collinsta heterophylla alone, b) Phaceltia
distans and Amsinckta intermedia, c) Collinstia and Phaceltia
d) Collinsia and Amsinekta. Six larvae were isolated in
each vial, and three additional larvae were added to c) and
d) after three days. In each case the only successful es-
tablishment occurred in flower buds of Collinsta, with about
33% of the individuals successful. The Amsinekiq did not
fare well under the conditions and was essentially wilted
within three days, while the Phacelia remained in good con-
eieven tor at least five days. All plants were in poor con-
dition by eight days, and the a) vial became diseased by the
llth day. There was no evidence that feeding occurred on
either Cotlinsia leaves or any part of the Phacelta or Am-
stnekta. Surviving larvae, along with others established in
separate containers with Collinsta bouquets were fed subse-
quently on flowers of Collinsta heterophylla which had been
taken in immature condition on March 25 at Raines Park and
kept in water, where the plants developed to full bloom.
Additional C, heterophylla from the Botanical Garden was
provided to nearly mature, 24-26 day old larvae.

Larvae of all instars fed within developing flowers. If
a bud was entered, no feeding occurred on sepals except in
excavating an entrance hole; ovaries, stamens, and corolla
parts were fed upon, preventing the bud from opening. Usu-
ally partially or fully opened flowers were used, and ovar-
ies were consumed, along with basal portions of the corolla,
but no feeding occurred on the sepals. After a few days
young larvae migrated to new flowers, leaving the wilted
corolla in sttu. In the field this resulted in several
withered and abandoned flowers on a given plant, indicating
the presence of one larva. Normally only one or two flowers
on any given tier were affected. The larva moved upward as
the inflorescence elongated, rather than working around the
inflorescence until all available flowers at one tier level
were exhausted.

In contrast to some species of Ethmta, the larvae curled
and feigned death at the slightest disturbance. They were
thus difficult to manipulate during transfer from one flower
to another, as they could not be induced to spin silk onto
the probe, and even if transferred with a damp brush or for-
ceps and balanced in an immobile posture on a new flower,
would almost always drop off upon moving again. However,
they showed a strong tendency to wander up vertical objects,
and usually migrated back up flower stems and reestablished
on their own when a given flower became exhausted.

Development proceded rapidly relative to other Ethmia.
Individuals provided with Colltnsta buds in good condition
reached the second instar by 8-10 days, and the third in-

Star by 11-13 days. The fourth instar was not preserved,
but all larvae had passed through it by 25-27 days. Final
instar, 25-29 day old larvae were preserved on April 25 and

27, and the last mature larva prepared for pupation on
April 29, 31 days following beginning of egg hatch.

16 seylla

On April 27 the Raines Park site was revisited and
larvae were found abundantly in Collinsia flowers (69D60).
Development was retarded relative to that in the laboratoty.
No larvae were preserved on this date, but after six days
storage at outdoor temperatures, a larval sample showed
three instars, 3rd, 4th, Sth, im apd: “4: 5:2 patie emer
days all three. were still present, in thie fatio se gaeeee
Only full grown larvae remained on May 9 (12 days after col-
lection), 51 days following the original collection of fe-
males in the field. Thus height of oviposition probably
occurred March 19-26, and most larvae reached maturity in
the field about May 3-9, an average lapse of 45 days.

There are five clear-cut larval instars, according to
unsexed head capsule measurements (fig 4).

First instar: Length 1.2-1.4 mm; HC 0.16-0.18 m,
light brown; ThSh narrowly light brown, well defined; body
yellowish, integument unpigmented.

Second tnstar: Length 2.0-2.5 mm; HC 0.25-0.27 im,
dark brown; ThSh well defined, brown; integument with pale
DL color, faintly ,efined paler D) and rings, around mae
Which are barely visible; AbCr 8-9, essentially a complete
eirele; AnCr 7-8.

Third instar: Length 3.2-4.8 mm; HC 0.38-0.43 mm,
dark brown, slightly mottled paler; ThSh brown, siiemmi
mottled darker posteriorly; D well defined; unpigmented ex-—
cept slight median pink line; ‘DL pink, weakly deztimed ial
pinkish, scarcely defined; Pin dark, well defined; Abdlxz
Ho1I8 Wao G9) - hs aN

Fourth instar: Length 5.5-7.2 mm; HC 0.51-0.68 mm,
light brownish mottled with darker brown; integument pattern
as in final instar, paler; D well defined, unpigmenteawer

cept median pinkish line; DU fairly well defined? eum aia

surrounded by unpigmented circles; AbdCr 10-14 (mostly 10-
a) eA Cte =O

Fifth instar: Length 8.8-12.0 mm; HC 0.82-0.98 mm,
orange with faint brown mottling; ThSh sclerotized as median
lateral spot and posterolateral blackish patch; integument
pattern well defined pink on whitish—inpigmented of pumpiatoa
on pinkish-unpigmented (probably varying with petal colors
consumed), D well defined with median pink streak; DL well
defined, dark, defining unpigmented circles around the black
Fin, L unpagmenveds; “iN fasiciiy ewe il die raned od er 14-18,
rarely 20, usually nearly uniordial: AnCr 162W/7e)lode eriene
secondary setae in LV group on abdominal Sseemenvs) Woy 2anuee
sometimes 8. [ae

Pupa. > Smallablocks of idnjyuvucea whrpoler sh iota
Stalks were provided and were used by all successfully pu-
pating individuals. Each spun the cocoon in a narrow gal-

lery running paral lely with 2thie igiraiin Jon) ihe wood. It

Biological studies on Ethmia 17

appeared that abandoned Cossonus gaileries were appropriated
and at times somewhat enlarged. Emergence trackways led out
to split ends of the substrate and each was divided into two
chambers by silken caps, one near the surface and one reces-
sed several mm, which was of slightly less diameter than the
pupation chamber, located at the anterior end of the cocoon.
Cocoons ranged about 5.4-5.7 x 1.3-1.7 mm and were simple,
without any interior meshwork.

Pupae (fig. 10) ranged 4.7-5.2 mm in length and were
simple, without functional cremaster, anal legs, or other
setation. The ninth segment was unmodified and fused to
the eighth at mid venter. The spiracles were small, simple,
0.35 mm in diameter. Evidently the cocoon shape retains the
pupa at emergence.

Pupae formed by larvae in May, 1969, failed to emerge,
although housed at Russell shed, where conditions stimulated
emergence of plagtobothrae (69D58) in the same winter. Pu-
pae were still viable appearing when extracted from cocoons
after 17 months.

ETHMIA BREVISTRIGA BREVISTRIGA CLARKE

Etpmea brevistriga Clarke, 1950, Jour. Wash. Acad. Sci.,
I@.=2k6.3).

The nominotypic subspecies is known only from locali-
ties along the immediate coast of California.

Study areas. - 1) Laguna Puerca, San Francisco; adults
common in association with Phaceltia distans, April 7, 1961;
5 males, 5 females retained alive (C.D. MacNeill and J. Pow-
Poy (6bp2): larvae’ on P. distane, May 6, 1961 (6102); lar-
waeian 2. distans, May 24,-1961 (61621). 2) Lobos Creek,
San Francisco; adults in association with P. dtstans, April
7, 1961; 1 male, 1 female retained alive (6/23

Adult behavior. - A single, well defined flight was
shown in 1960 and 1961 at San Francisco, from mid March to
mid April, prior to beginning of flowering of Phacelta dis-
tans. Six pairs were caged with a bouquet of P. distans.
These moths showed a slightly later diurnal activity period
than albitogata, housed under similar conditions. Individu-
als of brevistriga were not active before noon, and even by
1:30 P.M. only limited movement and no oviposition was occur-
ring. The height of activity appeared around 3:30 to “230.
continuing until about sunset, around 6:00 PiuM~ | By. (300 they
had become sluggish and only flew straight down if dislodged.
None moved at night. This species showed a greater tendency
to perch on the host plant than any of the others studied.
About 60-80% of the individuals rested on the plant, even at

night.

Mating was witnessed twice. A pair was swept tn copulo

18 brevistriga

at Lobos Creek at 3:00 P.M. They had been flying or were
perched onthe tip of "a Phacelva branch.” They Were) acest
coition at 7:00 P.M. following transport from the frend:
Housed in darkness, they remained in copulo until at least
11:00 P.M. The second pair mated sometime between 1:00 and
5:30 P.M. on the first day after confinement; after 5:30

they remained inactive, clinging to an upright pin al lonight.
Separation occurred between 9:30 and 10:10 A.M.

Oviposition by several females was observed, between
3:30 and 4:00 P.M. It probably took place earlier, and one
female exhibited apparent oviposition behavior at 5:15 P.M.
Characteristically females crawled over the uppermost foliage
or moved spirally up a stem, with the abdomen distended,
moving rather slowly and vibrating their antennae. The sub-
strate was tapped 4 or 5 times with the papillae anales pri-
or to deposition of an egg!” A. period of ‘quiescence mulomces
45 seconds) usually followed each egg after which the moth
resumed its crawling or flew to a new spot. Periods of
crawling on the screen were sometimes interspersed with those
of oviposition.

About 80 eggs were deposited by the females during the
first two days of confinement. It appeared that none were
laid on the plant after the third day. Nearly all were con-
centrated in the upper 5 cm of foliage, mostly around the

buds. The eggs on the inflorescences were not nested deeply
Into crevices, but were placed between the plant hairs (figs.
21-23). The uppersides of upper, young leaves and the sma

stem were also used as oviposition sites. Eggs placed below
the upper 5 cm of foliage were on the stem. None were
placed on the undersides of the leaves except on the main
midrib. \ Some oviposition after the third of foureh) davon
confinement took place on the cardboard floor of the jar.

Egg. - The eggs ranged 0:31 x 0.53 mm to 0.30) x 040mm
and as thick as wide ‘(fig. 23))2° When first depositecmeamer
were pearly white; after about 48 hours they turned yellow.
Prior to hatching the dark larval head capsule became visible.

Eggs hatched April 19-20, after about 11 days at room temp-
erature.
Larva. - Several Phacelta distans plants from Laguna

Puerca were planted in pots in a greenhouse prior to emer-
gence of the first instar larvae. These plants, which ma-
tured sooner than those in the field, were used for observa-
tions on behavior of young larvae.

First instar larvae migrated upward and commenced feed-
ing at bases of flower buds. In one instance a one day old
larva had bored through the sepals of a small unopened bud.
The first external evidence of established larvae appeared
by 4-6 days ‘in’ the form of ‘Small’ frass accumulations simeene
flower heads.

By the 14th day the insectary plants had bloomed com-

Biological studies on Ethmta 19

pletely, but the larvae, in the third and fourth instars, had
prevented development of some buds in which they fed. In
each case the larva had formed a well concealed shelter be-
tween the rows of flowers on the scorpioid spike, hidden
primarily by the dense plant hairs. The shelters were held

Hegerner ‘by a weak network of silk. . Feeding oe@curred in the
currently opened flowers and unopened buds, usually all the
Wayeoue to the terminal end. of .the inflorescenee. |Not all

of every flower was consumed, and some were still in apparent
bloom. Damage to the inflorescence was not evident external -
ly, and the frass accumulations were the only visible evi-
dence of the larvae.

On May 5, 17 days after commencement of hatching in
the laboratory, the Laguna Puerca site was investigated.
Three third instar larvae were found, in shelters similar to,

but less extensive than, those in the greenhouse. Field
plants had bloomed only about half way along the infloresc-
ence. Larvae were located just basad of the current bloom,

feeding on the flowers with developing seed. Frass from
these shelters was not visible from the exterior, evidently
having been dispersed by factors such as wind. All larvae
were moved to newly potted plants at this time, but the
plants did not survive. The remaining larvae were transfer-
red to salve tins with cut inflorescences two days later.

By the 24th or 25th day following hatching some larvae
fag freached the last instar. One larva on one of the origi-
nal potted plants reached the last instar by the 28th day,
when it was preserved.

A third examination of the field colony was made on May
24, 36 days after laboratory eggs began hatching and 6 days
after the first cocoon was spun in’the greenhouse. Larvae
were found to be fairly common in areas where the Phaceltia
was more sparse, although the shelters were as inconcpicuous
as they had been three weeks earlier. By this time field
larvae were mostly penultimate instar; a few were antepenul -
timate, and only one was in the final instar.

All larvae reached the final instar by the eighth day
after the second larval collection, and the final larvae
which had not spun cocoons were preserved June 6, 60 days
aimeer the original adult collection.

There appeared to be five instars, on the basis of un-
sexed head capsule measurements (fig. 3).

First instar: Length 1.5-1.7 mm; HC 0.18-0.20 mn,
brown; ThSh and anal shield pale brown; integument and setae
unpigmented.

Second instar: None preserved; HC 0.27-0.36 mm [5],
brown.

20 brevistriga

Third instar: Length 4.3-6.0 mm [3]; HC 0.42-0.47 mn,
dark brown; ThSh brown; Pin minute, dark; DL sometimes with

a trace of pale brownish, AbdCr 8-11; AnCr 9.

Fourth instar: Length 5.3-9.3 mm; HC 0.66-0.77 mn,
usually slightly to considerably paler brown than thigaan—
star, lightly mottled; ThSh paler brown; Pin small; integu-
mental pigment well developed, D white, DL brownish, broad,
extending below spiracle; L narrow, whitish; LV with Jitcle

pigment; AbdCr 9-13; AnCr 9-10.

Fifth instar: Length 9.0-13.2 mm; HC 0.86-0.95 aim,
orange brown, mottled; ThSh pale brown with dark spots; D
white (not unpigmented), DL broad, as in fourth instar,
darker, purplish; Pin small, in DL surrounded by Whitien
circles; L.white; AbdCm 157-18;, AnCr 16-17. Segment AQ with
6-8 tiny secondary setae on LV.

Pupa. - Pupation and successful development took place
in small beetle galleries in split sections of Yueca whtpplet
inflorescence stalk, in one instance about 15 mm into the
yucca, although not much excavation of the matrix by the
Ethmia larvae was involved. Pupation also occurred in a
corner of a salve tin, .in flower ,heads,, and in #0) dSMetea-
per toweling. The only successful emergence occurred from
one of the latter. Those in the flowers and salve tin des-
sicated prior to development.

The cocoon surface was papyrus-like, without much
loose internal silken mesh. The pupae ranged 5.4 to 5.6 mm
in length. The anal legs were short, the free portion only

0.22-0.23 mm long, appressed to abdomen, with 16-20 setae
Situated broadly over the distal end.

ETHMIA BREVISTRIGA ARDICOLA POWELL

Ethmta brevtetrtga ardicola Powell, 1971, U. Caligiy apes
HQ Jy) Le pisesisn.

This race occurs at inland stations, mostly in the
mountains marginal to the deserts. From the following frag-
mentary data and larval morphology, ardtcola appears to
have essentially the same biological characteristics as the
nominate subspecies.

Study areas. - 1) Hills two miles northeast of Lake-
side, San Diego Co.; adults taken in flight without definite
plant association, Mareh 13-,.°1963 /(WJuAe Chemsak” ands eee ome
ell); S males, 1 female retained ial ive (63C2) 2) 2) pian
Flat, 16 road miles southwest of Palm Desert, Riverside Go.
adults abundant at flowers of Cryptantha ?etrecumetssa and
Flying in association with Phacelta dtstans subsp. australts
April 7, 1963 (R.L: Langston, C.A. Toschi (and) J). Powe tie
females retained alive (63D6); April 12, 1963, 4 males 5 fe-

Biclogical studies on Ethmia 21

males retained alive (63D17); young larvae on P. distans var.
Bweeralts, April 13, 1963 (63D20).
Adult behavior. - This subspecies has about the same

seasonal flight period as its coastal counterpart, despite
the higher elevation of the inland sites (up to 5000 ft.).
Moths of 63C2 were caged in a gallon breeding jar in the
Field with a bouquet of Cryptantha and kept under variable
conditions until the fourth day. They did not begin activi-
ty until about 12:20-1:00 P.M. with the room temperature at
about 18° C, even though an Ethmia minuta male in the same
jar had been active for two hours. As with b. brevistriga,
the greatest activity seemed to be about 4:00-4:30 P.M. The
last individual ceased activity and entered the quiescent
posture at 5:40 on one afternoon, but several were active un-
til 6:10 (dusk) on another; and moths were observed with the
antennae in active position as late as 7:00 P.M. on the tenth
day after collection.

Males lived 8-13 days and the female 13 days, but only
2 eggs were deposited, those on the glass side of the con-
tainer. Presumably absence of Phacelta resulted in failure
to initiate oviposition. Moths of 63D17 were caged in an
85 x 100 mm jar with a bouquet of Phacelia and Cryptantha but
became water soaked in transit in an icebox from the field
laboratory April 14; several recovered and two females lived
until the sixth day following collection. Eggs were deposi -
ted April 14-17 on both upper and lower surfaces of Phacelia
leaves, not on buds, and on Cryptantha foliage, dry Cryptan-
tha flowers, and on a dead Ethmia male.

The three 63D6 females deposited 1, 6, and 10 eggs in
their individual, dry vials during the 2-3 days they lived.

Egg. - Eggs deposited in dry vials were evenly oval,
tapering slightly towards both ends, not as rectangulate as
in related species. The width and length ranged 0.33 x 0.63
fe 8 60x-0 563 mm.

Eggs of 63D6 were stored in dry vials in warm condi-
tions of a field laboratory and hatched in 8 to 9 days;
those of 63D17 were stored under moist conditions and vari-
able, cooler temperatures (including one to two days in a
field icebox) and hatched in 8 to 13 days.

Larva. - First instar larvae hatching from eggs on the
plant material, Phacelita dtstans subsp. australts and Cryp-
tantha citreumetissa were left in sttu in the inflorescences,
Most established feeding sites successfully on Phacelia buds,
although leaf material was eaten by two individuals. None
fed on the Cryptantha. Those from dry vials were placed on
flowers of Phacelia, and the flower parts served as food
throughout their growth.

The second instar was reached by about the sixth day
by most larvae; thereafter developmental rates varied, Owing

22 albitogata

to vartatton.tn GConditron of the olant, maternirar.

The Phacelta stems in one of two 63D17 lots began to
rot a week after the larvae hatched, and these larvae were
transferred to fresh, although mature, Phacelta dtstans from

Stanislaus County. They continued development, using mature
flowers; both the flower parts and developing ovules were
eaten. One larva reached the final instar by the 30th day.

A second lot was retained on the original Phacelta ma-
terial from Riverside County, which remained in good condi -
tion for about 27 days after the larvae hatched. However,
all flower parts were eaten by this time. Larvae fed entire-
ly on the half of the flowers towards the center of the
spike, or by cutting a hole through this side and eating
the center portions out, taking whole developing seed or
only their inner haif. By the 33rd day the plant had dried
excepting the stems, and larvae starved tn the final three
instars.

Laboratory reared larvae averaged somewhat smaller and
were considerably paler than b. brevistriga.

First tnetar: Length 1.3-1,4 mm, HC 0.19-0020"imeat
most colorless except ocellar area black.

Second tnstar: None preserved; HC 0.27-0.36 mm [3],
pale brownish.

Third instar: Length 3.6 mm [1]; HC 0.46-0.49 mm,
dark orange-brown; ThSh pale tan; integument, setae and
crotchets unpigmented.

Fourth tnstar: Length 6.4 mm [1]; HC 0.63-0.68-mm [4],
orange-brown, mottled; ThSh orange-brown; integument, setae
and crotehets unpipmenteds BbdCr 10-01. AnCreier

Fifth tnstar [2]: Length 7.5-8.0 mm; HC 0.79-0.85 mm,
orange, mottled; ThSh orange-brown; integument pattern simi-
lar to b. brevistrtga but much paler and reduced; D whitish
(not as distinctly white); DL dark pinkish or rosaceous; se-
tae and crotchets unpigmented; AbdCr 14-16; AnCr 14-16.

ETHMIA ALBITOGATA WALSINGHAM

Ethmia albitogata Walsingham, 1907, Proc. U. S. Natl. Mus.,

33% L209.

This species is known from only a few localities in
central California. It is closely related to F. plagtoboth-
rae, and it was not until differences in hostplants and lar-

vae were discovered that distinguishing morphological char-
acters in the adults of the two species were recognized.

Study areas. - 1) San Bruno Mountains, San Mateo, Co.;

Biological studies on Ethmia 23

adatts common, Feb. 28 - March 1, 1963 (C.D. MacNeill and J.
Powell); 5 males, 2 females retained alive (63B9); larvae on
ameenekta Lunaris, May 3, 1963 (63E1).. (2) Pt. Reyes, Marin
Ee.c Jatvae on Amstnckia -spectabitlis, April 31, 1967 (G.A.
Gorelick) (670149). 3) Arroyo Mocho, 15 miles southeast of
Livermore, Alameda Co.; adults common, Feb. 22-24, 1968 (C.
D. MacNeill and J. Powell); 5 males, 4 females retained alive
(68B178); negative results checking Amsinckia and Plagtobo-
weg April 27,1969; adults sparse, Feb. 21, 1970.

Adult. behavior. - The moths fly in early spring, late
January to early March. Presumably germination of the host
plant has begun, but | have been unable to locate young Am-
stnekta when the moths are flying. Adults (63B9) were caged
with a bouquet of Phacelta californica, a suspect host;
while 1968 adults were provided with young Plagiobothrys no-
tnopulous, OWing to a misidentification of the moth. . The
correct foodplant was never offered as an oviposition sub-
strate and stimulus.

The moths exhibited a definite diurnal activity rhythm,
but neither mating nor oviposition was observed. Individuals
from the San Bruno Mountains commenced activity earlier, be-
ginning to abandon the quiescent posture by about 9:30 A.M.,
three hours after daybreak, with the outside temperature at
9-11° C. During the following two hours all adults engaged
in some movement. One female was observed to take water in
this mMatinal” "pre-activity'’ period. Continuous activity,
with moths mostly crawling at the side of the jar towards
the sunlight, took place between 12:00 and 4:00 P.M. By
h:30 some moths ceased movement, and by 5:30, with the last
rays of sunlight on the jar, most individuals had assumed
the quiescent posture. Adults from Arroyo Mocho, by contrast
remained inactive until 10:30-11:30 A.M., even when the con-
Eamher was transported by car 15 miles to Berkeley. How-
eve they remained active later, till 6:00-7:00 P.M.,
through the dusk period. Outside air temperatures were warm-
er during the 1968 observations, ranging to 24° C maximum
compared to 15-17° in 1963. Moths in both groups generally
displayed a longer activity period than some of the other
diurnal Ethmia, and they became active quickly if exposed to
direct light at night.

Males lived 2-6 days, females 6-9 days, following col-
Pection “in the field.

In 1963 no eggs were deposited by captive albitogata.
Presumably the Phacelia did not provide adequate stimulus,
since cage conditions were comparable to those extant during
successful oviposition by other diurnal Ethmia. The 1968
females: laid only 35 eggs; again absence of Amsinekta prob-
ably adversely affected oviposition behavior. Of the total,
28 eggs were placed on the helical, ribbed portion of a hor-
izontal, screw-cap vial which held a moisture wick and
Plagiobothrys bouquet. The remaining eggs were deposited
on Plagtobothrys leaves (5 upperside, 2 underside).

24 albitogata

Egg. =) The eggs didjinot differ super f cnc lily aie
those of plagtobothrae. No measurements or photographs were
executed... |incubation required iol) 25 day s,.

Larva. - First instar larvae were placed on Plagto-
thrys nothofulvus from Arroyo Mocho. Uprooted young plants
had been placed in water vials 12 days previously, and after
initial die back of lateral leaves, survived well for the
duration of the experiment and were sending up floral stalks
by the time of larval hatch. Most larvae did not establish
OM) seh is) shiosits lt appeared that those situated on new term-
inal leaves were unable to penetrate the thick pubescence;
when transferred to the undersides of basal green leaves,
where pubescence was less dense, most still did not feed
successfully. Two individuals accepted the Plagtobothrys
and fed about 7-9 days, reaching the second and third instar.
Feeding occurred in the form of small skeletonized areas,
with a thin silk envelope between the new leaves of the ter-
minal growth. Although the plants remained in good condi-
tion more than three weeks after transferral to the water
vials, no flowers had begun to open by the time the larvae
succumbed in what appeared to be starved condition. The un-
availability of flower parts rather than the wrong host ge-
nus may have been the critical factor in the failure of the
larvae to mature.

At San Bruno:Mountains in 1963 close association of
adult flight with several clumps of poison oak (Rhus diver-
siloba) on rocky outcroppings, enabled discovery of larvae
on Amsitneckia lunarts there in early May. At this time, about
60 days after the height of observed adult activity, the
plants were in full bloom, and larvae of the final three in-
stars were present. There was variation between Amsitnekia
colonies from 75% penultimate and none full grown to 50% ma-
ture final instar and the remainder young final instar.

All larvae were found inside inconspicuous shelters
formed in the flower spikes, similar to those of £. brevt-
strtga. Larvae moved along the upper side of the scorpiod
spike, webbing the flower parts together above the larval
galleries. Feeding took place on the whole inner side of
the flowers. Developing ovules and ovaries of unopened flow-
ers were consumed, and even sepals were eaten by larger lar-
vae. Portions of the flowers on the outer half, visible
from the exterior of the inflorescence, were untouched. Pet-
als were mostly above the area of feeding and remained in-
tact, without discoloration or wilting. Some feeding occur-
red on the inflorescence stem; in one instance. it Was) vomit
entirely through.

Shelters were not evident from the exterior, but affec-
ted inflorescences could be detected by frass clinging to
the older, unoccupied portion of the spike where elongation
of the stem caused separation of the partially eaten flowers,
exposing silk. Frass apparently was entirely retained with-
in active parts of the shelter, not visible from the exterior.

Biological studies on Ethmia 25

Larvae were placed in plastic sandwich boxes with small
bouquets of Amsinekia. However, the viscid plant did not
keep well and mould developed within three days. Yueeq
pith was added as a pupation site, but the colonies became
affected with disease and no larvae pupated.

As in plagtobothrae, head capsule measurements did not
enable definition of all instars (fig. 2). In both species
either the second instar head capsules were not recovered or
a greater relative size increment occurred between the
first and second than in other Ethmia. Moreover, these two
species exhibited two color phases in the final instar, a
characteristic not observed in related species. Head cap-
sule measurements showed only a very slightly larger average
in the paler of the two phases.

First instar: (None preserved in healthy condition)
HC 0.18-0.22 mm, dark brown; integument, including pinacula,
unpigmented.

Seeond instar(?): [1] (Not preserved in distended con-
dition) HC 0.33 mm; dark brown; integument unpigmented, Pin
Slightly darker; AbdCr 9-10; AnCr 9.

Third instar (?): [i] (Not preserved in distended con-
dition) HC 0.50 mm; brown, unmottled; ThSh brown laterally,
unpigmented mesally; D whitish, DL brownish gray, L unpig-
mented, LV pale grayish, Pin dark, not defined by pale ar-
eas; AbdCr 9-10; AnCr 8.

Penultimate instar: [4] Length 7.5-7.8 mm; HC 0.57-
0.60 mm, dark brown, paler above labrum, not mottled; D
searcely distinguishable, whitish, DL pale grayish (not dis-
tinet as in fourth instar plagiobothrae), L unpigmented;
AbdCr 6-9 (usually 8-9); AnCr 6-7.

Pinal instar: Length 10.2-13.0 mm (rarely, teneral?,
7.7 mm with integumental pigment reduced); HC 0.83-0.96 mm,
strongly mottled; similar to plagiobothrae but paler, with
the two color forms not as distinguished: (a) (HC avg. 0.88
mm) D pale without yellow-orange spots, DL well defined,
dark to pale gray; (b) (HC avg. 0.90 mm) D pale with segmen-
tal yellow-orange spots, DL and LV pale gray; Pin small,
well defined, dark; AbdCr 10-20 (usually 12-16); AnCr 12-16.

ETHMIA PLAGIOBOTHRAE POWELL

Ethmia plagtobothrae Powell, 1971, U. Calif. Publ. Ent.;
in press.

Although discovered only about ten years ago, this
species has been collected many times. The larvae are often
encountered in large numbers, but in the laboratory they are
extremely susceptible to disease. Those which pupate fre-
quently do not metamorphose. Only a few adults have been

26 plagtobothrae

taken in the field. Ethmta plagtobothrae is closely allied
to albitogata, but the two exhibit marked biological «diftfter-
ences.

Study areas. - 1) Cool, El Dorado Co.; larvae abundant
on Plagtobothrys nothofulvus, April 24, 1961 (C.D. MacNeill
and J: Powell) (6104, °61D5); 1 male,-4 females Manchmeue
1962, female retained alive (62C2); investigated for adults,
March 29, 1964 and March 23, 1965, negative results. 2)

Six miles west of Whiskeytown, Shasta Co.; larvae on Plagto-
bothrys probably nothofulvus, May 10, 1961 (R.L. Langston
and J. Powell) (61E10). 3) Eight miles south of Leesville,
Colusa Co.; larvae’on P. nothofulvus ‘April 12,01962 .aaoee
Chemsak and J. Powell (62D5); investigated for adults, March
8, 1963, negative. 4) Elk Mountain, 2800-3000 feet, 11-12
miles north of Upper Lake, Lake Co.; 3 females flying in
association with, 1] egg, 2 young larvae on P. nothofulvus,
April 4, 1962) (62D12);- investigated for adults, Aprimmas
1964, negative (penultimate instar larvae on P. nothofulvus
at 2200 feet, one mile south on the same road); adults
sparse, March! 18; 1965 .(R-&. Langston and J. Powe ki )emeeg
Ten miles south of Creston, San Luis Obispo Co.; 3 larvae on
P. nothofulvus, April 30, 1962. 6) Arroyo Mocho, 16 antes
south of Livermore, Alameda Co.; larvae on P. nothofulvus,
April 30, 1963; investigated for adults, February 23, 1964,
March 14, 1965, Feb. 24, 1968, March 4, 1969, Feb.’ 21, 90970)
negative. 7) San Antonio Valley Ranger Station, Santa Clara
Co.; mature larvae on P. nothofulvus, April 30, 1963; inves-
tigated for adults, Feb. 9, 1964, negative; 3 young larvae
on P. nothofulvus, March 14, 1965; ‘investigated Feb. 24,
1968, negative; 1 male, 1 female, March 4, 1969 (P.A. Opler
and J. Powell); larvae common on P. nothofulvus, April 27,
1969 (69058). 8) One mile north of Posey, Tulare Co; larvae
common on P. nothofulvus, May 14, 1963 (C.A. Toschi and J.
Powell) (63E2). 9) Three miles north of Havi bah, Keen Gee,

larvae on P. tenellus, May 15, 1963 (63E10). 10) eae
miles southwest of Havilah, Kern Co.; larvae on P. nothoful-
vus, May 15, 191637. 11) Havilah, Kern Co.; larvae common on
P. nothofulvus, April 28, 1964 (C.A. Toschi and J. Powell)
(64010). 12) One mile east of Woody, Kern Co.; larvaeuon
Plagtobothrys, April 25, 1964 (C.A. Toschi), May 3, 1964
(64016). 13) Three miles northwest of Mariposa, Mariposa
Co.; 3 young larvae on Plagiobothrys, March 25, 1965.

Adult behavior. - This species has a single annual
flight period, in early spring. The few field collections
indicate the moths fly in March and early April, ahead of or
at about the time the blossom period of the host begins.

The development of Plagtobothrys is highly variable from one
season to another at a given locality, possibly correlated
with early spring rainfall, and occurrence of young larvae
in mid March, 1965, suggests that the moths are sometimes
flying by mid February.

Only one female was observed in captivity (62C2).
This field collected individual was caged in a one-gallon

Biological studies on#thmia 27

Jar with a flat of planted herbs from Cool, and a bouquet of
Phaceltia caltfornica from San Mateo County. The female show-
ed the same activity periods and quiescent posture at night
as described above for £, albitogata. On one occasion, the
female did not move when lights were intermittently on from
dusk until 8:00 A.M. In this instance she began activity
about 10:30 A.M.

Oviposition behavior, with the abdomen distended and
curled towards the leaf substrate, was observed at 12:15,
2:00 and 4:55 P.M. one day, and deposition of eggs was wit-
nessed at 2:30 on another afternoon. On one occasion two
eggs were deposited in rapid succession, (a few seconds in-
terval) without apparent probing of the ovipositor, followed
by a third egg nearby a few minutes later. A total of 62
eggs was recorded. About two-thirds (44) were deposited on
the Phacelia (figs. 24-25) and 5 more on grass blades and
the glass side of the container adjacent to the Phacelia.
The remainder were located on what was presumed to be basal
rosettes of young Plagtobothrys. All but one of the latter
group and 90% of those on the Phacelia were placed on the
undersides of the leaves, which were more hirsute in both
cases.

The moth lived nine days after its capture, but prob-
ably no oviposition took place after the fourth or fifth day.

Egg. - (Figs. 24-25) The eggs were more variable in
shape than most other Ethmia studied, ranging in outline
from rectangular-oval to ovoid, tapering at both ends; width
and length varied accordingly, from 0.22 x 0.47 mm to 0.28
x 0.42 mm.

Development at laboratory temperature required 10-11
days, hatching April 2-4, 11 to 13 days after the female was
first caged. Some of the eggs were stored in a refrigerator
for 72 hours, and emergence of these was delayed about 3
days beyond the last of the non-refrigerated ones.

Emergence frequently took place off center from the mi-
cropyle, by means of an irregular slot contrasted to the
more or less evenly oval hole in the middle of the micropy-
lar end, which is usually cut by larvae of other Ethmia.

Larva. - Newly hatched larvae were placed on Phacelia
ecaltfornica, but none successfully established themselves.
A few small spots of skeletonizing represented the only feed-
ing and none of these larvae reached the second instar.

Two young larvae were collected at Elk Mtn., in the
basal rosettes of Plagtobothrys, but their shelter and feed-
ing were not observed. When preserved, they were in the
second and third instar.

Larvae of at least the final two instars fed entirely
exposed, on the flowering stalks of Plagtobothrys. No visi-
ble silk nor other shelter was employed. Larvae could be

28 plagtobothrae

found curled around the uppermost flower, feeding on the in-
florescences.

Frass was flipped free of the flower parts, appearing
on the sides of the container.

In the following larval diagnosis, specimens from Cool
and Havilah are mixed in the final two instars. Those from
Havilah average smaller, but the ranges of variation are
similar.

Evidently this species has five instars (fig. 1), with
two well defined color phases in the final instar. It was
originally assumed that two instars were involved, but head
capsule measurements do not show an appreciable separation.
Form (a) has dark gray, almost black integumental markings
with yellowish spots in the dorsal band, whiie (b) has much
paler gray integumental bands and conspicuous orange blotch-

es on each segment except the prothoracic. The pale form
(b), larvae seemed bulkier and apparently were more mature,
but | had no evidence that an ecdysis occurred in develop-

ments of (a)utonn bye

First instar[4]: Length 1.0-1.2 mm; HC 0.20-0.22 mn,
pale orange with ocellar area black; integument and setae
colorless.

Second instar[1]: Length 3.0 mm; HC 0.36 mm, dark
brown; ThSh, Pin small, and setae dark, integument unpig—
mented; AbdCr 6-7; AnCr 7-8.

Third instarl1]: Length 3.3 mm; HC 0.49 mm, lighter
brownish; ThSh not defined; Pin»sbrown, as smali as) iaeead
instar; integument unpigmented; AbdCr 9-10; AnCr 11.

Fourth instar: Length 6.3-8.8 mm; HC 0.53-0.60 mm,
dark brown, poorly defined- pale area. above labrum; )22agemee,
dark; DL pale gray, D well defined, LV scarcely p»enem@eee.
AbdCr 9-11 (Cool) or 7-8 (Havilah); AnCr 9-11.

Fifth instar: Length, form (a) 10.0-13.0 mm, form (b)
12.9-15.8 mm; HC 0.75=0.85 mm (Havilah, (a) average Oni77,
(b) average 0. 79 mm), 0.77-0.88 mm (Cool, both forms average
0; 834mm), orange-brown, strongly mottled; ThSh darkened lat-
era iis, only; Panviblack< TD whare ewe defined, each segment
except prothoracic with a bright orange blotch as broad as
D; DL gray, narrow; L pale with dull, irregular blotch above
Spiracle; LV pale gray, irregularly mottled; AbdCr 16-20
(usually 17-19), biordinal mesally; AnCr 19-21, biordinal.

Pupa. -* Successful. pupation: occurred in| folds of@pager
toweling or tissue paper, and in yucca pith. Under labora-
tory conditions most individuals either died as prepupae or
young pupae or remained in diapause and did not emerge. On

two occasions (61D4, 63E2) single moths emerged early the
Following year and twice pupae were still healthy appearing

Biological studies on Ethmia 29

during the second winter: two pupae of 61E10 in December 1962
(19 months after pupating), and one of 64D10 in February

1966 (after 21 months). The latter was placed in an outdoor
cage at Berkeley through the spring, 1966, but still did not
emerge. Larvae from several lots constructed cocoons in
corners of salve tins, and in one case between a cotton plug
and glass side of a vial. In all these cases prepupal lar-
vae or pupae became dessicated and collapsed prior to devel-
Opment.

Full grown larvae from San Antonio Vailey (69D58) were
placed in 35 mm square pill boxes or small salve tins, two
or three individuals per container, with a block of yucca
cortex in each. After storage at laboratory temperature for
10 weeks they were transferred to the outdoor shed in July.
Successful emergence occurred by late February in over 60%
of the individuals, suggesting that temperature and moisture
rather than photoperiod are stimuli which are important to
development during the pupal stage. Cocoons were formed in
cracks in the yucca or between the yucca and paper liner.

Pupae from Cool ranged 5.0-5.6 mm in length. The anal
legs were dorsoventrally flattened more than in other Ethmia
studied. In addition they showed a definite tendency for
greater lateral expansion distally (appearing boot-shaped
in outline rather than evenly expanded laterad and mesad).
The free portion was about 0.24-0.27 mm long, with a lateral
projection of 0.09 mm. The legs had 32-36 (rarely 38) hook-
ed setae which are about 0.05 mm in length. The setae of
the cremaster area were 0.12 mm long and relatively strong,
remaining intact during emergence of the moth.

Natural enemies. - About 40% of the groups of larvae
in various types of containers became diseased and nearly
all larvae in these lots succumbed prior to pupation. Rep-

resentatives from two affected collections (64D10, 64D16)
were submitted to the Division of Invertebrate Pathology at
the University of California, Berkeley. G.M. Thomas respon-
ded (in 1itt.), indicating that media inoculated directly
from titurated specimens produced pure cultures of a Pseudo-
monas sp. and that observations indicated this bacterium was
the cause of the disease.

lt is assumed that the epidemics were brought on by
conditions in rearing, since similarly affected larvae were
not commonly seen in the field, and in at least two cases
(61D4, 64D10) containers with few larvae did not show the
symptoms while those with larger groups did. However, the
high incidence of these epidemics and the fact that such
symptoms occurred only in this species, albttogata, and
charybdis, indicate that the body flora of these larvae
differs from that of most Ethmia, causing them to be more
subject to disease. This may help account for the fact that
larvae of these three species are more easily found in high
numerical density in the field than the adults, whereas the
reverse is true with other Ethmia | have studied.

30 minuta

ETHMIA MINUTA POWELL
Ethmia minuta Powell, L971, U. ‘Calif. Publ. Ents, an geece.,

This species was collected in southern California as
early as 1916, but probably it was not recognized as an Eth-
mta owing to the small size. The elongated, strongly scler-
otized ovipositor and smooth egg are features unique to this
species among known New World E£thmia.

Study areas. - 1) Hills 2 miles northeast of Lakeside,
San Diego Co.; adults at midday flying and on flowers of
Cryptantha intermedia, March 30, 1961 and March 13, 1963.
2) Two miles northeast of Moreno, Riverside Co.; males fly-
ing in midafternoon, April 5, 1963 (C.A. Toschi and J. Pow-
ell); both sexes flying in association with Cryptantha tn-
termedia, April 12, 2:00-4:00 P.M., 4 males, 4 females were
retained alive (63D18).

Adult behavior. - Ethmia minuta has a single, early
spring flight, from mid March to late April. The moths are
diurnal. The four pairs from Moreno were caged in an 85 x
100 mm jar with a bouquet of Cryptantha April 13-14 under
field laboratory conditions. About 20 eggs were deposited
during this time, but no observations on behavior were made.
On the following day the moths became water soaked during
transport from the field in an ice box. Three females par-
tially or fubhly recovered, andsone Vived until: April tyme
was observed on the Cryptantha once, but no oviposition
occurred after April 14.

Eggs on April 13 were all deposited between bases of
flower buds (figs. 18-20). These were located only in in-
florescences with partially developed flowers. Those with
larger green seed and no blossoms left were not used for
oviposition. Evidently the elongated ovipositor of FE, mi-
nuta is an adaptation for use of the densely bristled inflor-
escences of Cryptantha tntermedta.

Egg. - (Figs. 18-20) In contrast to all other species
for which eggs were studied, those of minuta had a smooth
chorion, without visible network of structural ridges under
54x magnification. The shape was roughly oval, circular in
cross section, measuring 0.30 x 0.43 mm to 0.25 x 0.47 mm;
variation resulted from the situation of placement. Upon
dissection of the flowers most eggs were found to be wedged
between a sepal and upper portion of a carpel. Emergence of
larvae invariably occurred from the inward end of the egg,
adjacent to the carpel. Hatching occurred April 22 (# 10
days incubation).

Larvae. - Some of the buds on which the eggs were de-
posited had dried by the time the larvae began emerging, and
they were placed on the exterior of green buds. They seemed
unable to crawl on or penetrate the densely bristled vesti-
ture, and several died. None attempted to feed on stems;

Biological studies on Ethmta 31

no leaves were available.

By opening buds slightly with forceps and inserting
two day old larvae, |! was able to provide conditions which
enabled feeding. Whether the few larvae which successfully
established included any of these, or were only those which
entered directly from the eggs, was not determined. Feeding
by first instar larvae took place at the sides of developing
ovules. There was no feeding on petals or sepals, and lar-
vae placed in buds which were too young to have developing
ovules did not feed. Most died without establishing success-
fully, even after some feeding.

Those surviving hollowed out developing ovaries. Sec-
ond instar larvae were transferred on the fourth day from
the drying buds to buds which had been refrigerated two
weeks. Again larvae had to be placed into forced open buds
and not all succeeded in feeding.

On the twelfth day two third instar larvae were placed
on new Cryptantha from the refrigerator (originally taken
with the moths 21 days earlier). These larvae experienced
considerable difficulty and were unable to penetrate the
bristled buds after one hour. The two were then placed on
flowers which | broke open, but they were still unable to
Sstashish easily. -After another hour one larva succeeded in
beginning feeding on the inner side of the ovary wall. This
larva lived until the 17th day after hatching. By this time,
however, the 26 day old Cryptantha failed to take up water
when removed from the refrigerator and no additional food-
plant was provided.

First instar: Length 1.0-1.25 mm, HC 0.16-0.17 mn,
pale tan, almost colorless, ocellar area black; integument
and setae colorless.

Seeond instar: None preserved; HC 0.27-0.29 mm [2];
integumental markings evident as pale yellow-orange or
ochreous-tan blotches surrounding the DL pinacula.

Third instar[2]: Length 3.7-4.2 mm; HC 0.36-0.41 mn,
pale to dark brown; DL mottled pale olive-brown; Pin not
differentiated, setae and crotchets colorless; AbdCr +8;
AnCr +8.

ETHMIA CHARYBDIS POWELL
Ethmta charybdis Powell, 1971, U. Calif. Publ. Ent.; in press.

This bizarre species is known from only three locali-
ties, having been discovered when we reared a male from lar-

vae collected in 1967. The moth is unique among all EZthmia
by possession of extremely elongate and peculiarly thin legs,
as well as by markedly reduced mouthparts. Primarily on the

basis of male genital characters charybdts had been placed

32 Ccharypdts

as a monobasic species group related to the diurnal group.
However, with the discovery, in 1970, of the female which is
brachypterous, reevaluation of the species‘ assignment to
Ethmta is anticipated.

According to the geographical distributions of the
hostplant and of other Insects which occur at’ ther typemoca |=
ity, we expected colonies of charybdts along the western
edge of the San Joaquin Valley and in the Mojave Desert.
This has proven to be the case, with collection of larvae in
the Mojave in 1970 and recognition of apparently conspecific
larvae in the U.S. National Museum collection which had, been
taken in the southeast corner of San Luis Obispo Coun in
April 1956, ''sweeping wheat and various flowers", by G. Bee-
vor of the California State Department of Agriculture. Ex-
amination of Amstnekia tessellata in a similar habitat to
the type locality, at Jocalitos Canyon near Coalinga, Fresno
County, in early February and late March proved negative.

Study aneiash a ll) Big Panoche Creek, near Fresno-San
Benito County line; larvae on Amstnekia tessellata, April
20-21;°1967 (J. Powell’ and P.A. Rude) (67087) 2°) ‘vYounamterve
on Amsitnekta, March, 1968; negative results checking Am-
sitnekia tntermedia and A. gloriosa, March 5, 1969, and A. ~
tessellata, April 23, 1969, and with Amsinekta not yet germ-
inated, Feb. 5, 1970. ° 2) Ryan Mountain, Joshua Tree mnanenen-
al Monument; larvae on Amstnekia intermedta, March 31, 1970
(RAE. “Dietz eanded. Powells) © G7 0C 13) =

Adult behavior. - Only two adults, from Ryan Mountain,
have been observed alive, serving as indicators of the sea-
sonal and diel activity periods. Emergence occurred at the
end of November and beginning of December, after pupal
aestivation in closed containers which were housed under lab-
oratory conditions through the summer and in a modified out-
door situation during fall. Field surveys have not been
carried out during the fall months, but the normal flight
period is presumed to be late fall or winter, in part through
comparison with the life cycle of #. ttmberlaket, discussed
betow, and in part owing to the brachypterous condition of
the female in charybdts, a characteristic of certain winter
moths in other taxa.

As is true in other fall flying Ethmta, adults of ehar-
ybdits possess large eyes and nocturnal habits despite the
fact that activity, in November in desert habitats, must
take place in cold temperatures. Moreover, there was an in-
dication in laboratory charybdts that activity is restricted
to early morning hours rather than at dusk or early darkness,
when it was warmer. No crepuscular movement occurred, and
on several evenings activity by one or both individuals did

1 Probably in the vicinity of Cuyama “according vo Mina Gamqa—
ner, Bureau of Bntomology,) sacramenvo, (Californias.

Biological studies on Ethmia 33

not begin prior to 4-6 hours after nightfall. In every ob-
served nocturnal cycle both moths moved after 11:00 P.M.
This behavior may have been artificially induced, because
the moths were housed at about 16-18° C during daylight and
early evening, and in temperatures declining to about 12-14°
C between 11:00 P.M. and 8:00 A.M. It may be that optimal
temperatures for charybdis are well below 16-18°, and in the
field that might occur in early evening. ft ts HOt Unreas—
onable to suppose that this species is active at colder
temperatures than any other known Ethmta. By comparison,
the geometrid winter moth, Operophtera brumata(L.), in

which the female is brachypterous, has been observed to mate
and oviposit at temperatures just under 0° C (Cuming, 1961),
whereas most other Geometridae, even species which fly only
in early spring, are rarely active below a temperature

range around 4-5° C, and not at all below 2-3° C according
Eeutlight attraction records (Powell, 1962).

As in the case of ttmberlaket, Ethmia charybdis was

able to survive without water. The proboscis is short and
may not be functional. No moisture was provided during the
first 7-9 days the male was alive or the first 3-4 days foll-
Owing emergence of the female. Whereas a shorter period

would have been lethal to most Ethmta, there was no evidence
of weakening of the charybdts adults, and mating took place

during this time. Ultimately the male was killed when 10

or 11 days old, while the female, after an oviposition per-

iod during which she was provided with water, died 6-7 days

after emergence.

Mating occurred during the second or third night foll-
owing emergence of the female, when the male was 6-8 days
ola. sCopulation was initiated after 11:30 P.M., on an eve-
ning when both individuals had been active between 6:30 and
11:30. The pair remained in coition approximately 22-26
hours, showing no signs of activity during this time (even
when exposed to electronic flash and direct sunlight for
several minutes during photography). Separation occurred,
with both male and female moving away, between 4:00 A.M.
and 6-00 A.M.

Oviposition behavior was not observed, but it occurred
between 12-72 hours after completion of mating and could
have begun immediately in the matinal period following ma-
ting. Although no protein was provided, the single female
deposited 75 eggs, the highest total |! recorded for an indi-
vidual Ethmta. The eggs were placed in depressions and
holes in yucca pith and under and between layers of tissue
paper. The female selected cracks and open beetle galler-
ies In the yucca piece for 11 eggs, which were recessed up
to 0.6 mm below the surface. Most of the oviposition (58
eggs) occurred in creases, between layers, and onto the un-
derside of tissue paper liners of the original larval con-
tainers and fresh paper provided in the breeding cage.
Three eggs were nested adjacent to mouldy frass on the tis-
sue, suggesting that larval evidences on old foodplant may

34 charybdis

elicit oviposition stimulus. Dry flower and leaf fragments
of Amsinekia tessellata from a herbarium sheet were provided
but were not selected by the female.

Egg. - The eggs were characteristic of other Hthmza in
shape and chorion sculpture but were smaller than those of
other nocturnal species with forewing length comparable to
that of male charybdis. Eggs ranged 0.37 x 0.62 mm to 0.30
x 0.67 mm in outline, about the size of those of diurnal
species, which the female approximates in body size. Stored
at laboratory temperatures, all but a few apparently infer-
tile ones turned yellowish within 3 days, to a peach color
by 7-9 days, and later gradually reddish. They showed no
signs of maturation by the 15th day, suggesting diapause,
but they did not rapidly change to a tomato red color char-
acteristic of eggs in diapause in Ethmia ttmberlaket and in
tortricine moths (Powell, 1964). However, color transition
in ttmberlaket was not observed and may be a gradual, slower
process as in the present species.

Larva. - Younger instars were not observed; individuals
thought to be antepenultimate and penultimate were taken by
net sweeping. Mature larvae lived exposed on the inflor-
escences, usually perching on one side, below the highest
part of the plant, without any visible webbing. In the lab-
oratory those of the last two instars housed in 32 x 90 mm
plastic vials proved to be susceptible to disease epidemics,
even though individuals were separated, a few in each con-
tainer. Others placed in 25 x 75 mm salve tins were less
susceptible, and several matured successfully. Shelters were
spun among flowers, but these may have been constructed only
in preparation for pupation. Feeding probably occurs primar-
ily on deveioping seed and flower parts under natural condi-
tion's

At maturity larvae spun opaque cocoons in corners of the
rearing containers or in foliage (67D87). No soft, woody
substrates were provided. The 1970 larvae were offered yucca
blocks bearing galleries of cossonid beetles, but the three
larvae which successfully completed cocoons all selected
folds of tissue paper.

Penultimate instar (?) [2]: Length 9.7-10.0 mm. HC 0.58-
0.71 mm; orange, indistinctly mottled with brownish; ThSh
brownish, fairly well defined; integument color as in final
instar, paler than most but not all individuals; AbdCr 7-10;
AnCr 9-15.

Final instar: (Fig. 7) Length 10.0-16.0 mm. HC 0.85-

0.97 mm; orange, distinctly mottled with brown; ThSh not well
defined, sclerotized areas restricted to posterior margin;
integument pale to distinctly colored, D white with a median
rust-orange streak (sometimes reduced to a trace), DL fairly
uniform dark to pale gray, mottled, defining distinct white
circles around pinacula; L white, well defined, with variable
elongate blotch of pale to bright rust-orange, LV as in DL,

Biological studies on Ethmia oye

Bevaltiy slightly paler; Pin large, black, distinct; AbdCr
15-21; AnCr 21-22. Segment AY with O or 1 very small secon-
dary seta at LV, anal leg with small patch of usually 3 tiny
secondary setae.

pupae ~ Pupation occurred tn Various cocoon situations ;
but lack of suitable substrates may have ultimately resulted
in dessication of several prior to development. The only
successful emergences occurred from a cocoon tightly spun in
old foliage and flower parts in 1967 and in tight folds of
tissue paper in 1970.

Preserved pupae measured 6.0-6.8 mm in length, were
smooth, pale orange, without specialized spiracle structures.
The anal legs were rather short with slight to no lateral
extension distally, with 20-22 anchoring setae. The frail
cremaster homologue setae were located on a roughened, but
not depressed area.

ETHMIA ALBISTRIGELLA (WALSINGHAM)

Peeeqata albistrigetta Walsingham, 1880, Proc. Zool. Soc.
Remar, 1680789.

Described from the Siskiyou Mountains on the northern
border of California, albtstrtgella is widespread in western
North America, occupying more boreal regions than the close-
ly related nadia, discussed below. Both have small eyes,
but according to phenetic assessment are more closely rela-
ted to members of the semilugens group which are nocturnal,
than they are to the foregoing diurnal species (Powell, 1971).
lt is possible that small eyes and diurnal habits in albts-
trtgella are a secondary development as a result of adapta-
tion to high elevations where temperatures early in the sea-
son deter night time activity.

Seudy areas. - I) Chipmunk Flat, 3 miles’ west of Sonora
Pass, 00000 feet, Tuolumne Co.; adults associated with Pha-
celta ramosissima, June 25, 1962 (C.D. MacNeill and J. Powell)
20mates, 3 females retained alive (62F11)3; and July 1, 1962,

male, 1 female retained alive (62G4). .2) Donner Pass,
7300 feet, Nevada Co.; 1 male, 1 female flying near P. ramo-
stsstma, July 4, 1962 (C.A. Toschi and J. Powell); larvae

on P. ramosissitma. August 3, 1962 (62H3).

Adult behavior. - Ethmita albtstrigella has a single
annual generation, flying early in the season, in California
from mid June to. mid July, after the snow ‘In the vicinity has
receded to patches and the Phacelta has not yet begun to
bloom. Adults were apparently actively flying between 1:00
and 3:00 P.M., and [| have seen no record of collection of this
Species, at. Wight). In the breeding jar, adults were caged with
a bouquet of Phacelita from the collection site. They did not
show a definite activity period, but laboratory conditions
probably differed more (especially warmer in late afternoon,
night and early morning) from field conditions with this

36 albistrigella

boreal species than for any other species studied. Most
albistrigella moved occasionally or were actively crawling
during morning hours, as early as 8:30. Most were active,
crawling towards the daylight side of the container, feeding
at water, etc. between noon and 5:00 P.M. sHowevens wmane un
stances were noted in which moths moved after dark, both
with and without lights in. the room... They did not meen eto
be continuously active after dusk, but they were not consis-
tently inactive as in the cases of the diurnal species dis-
cussed above.

One mating pair was observed at 8:00 P.M., just prior
to sunset, having coupled after 6:30 P.M. Separation occur-
red, Ln the .darky@hetweenm 92 5 nancy lls O0n kM.

Oviposition was not witnessed, but took place between
1:00 and 2:30 P.M., and at least once between 8230 cme
(dusk) and 8:30 A.M., after six days confinement of the fe-
male. Eggs were placed only on the Phacelia leaves, except-
ing one or two placed on the glass adjacent to the Phacelia.
About 70 eggs were deposited by 3 females, almost exclusive-
ly on the undersides of the leaves; 2 each were placed on
Uppegsides of leaves and Jonisimens..

Egg. - The shape of eggs was more variable than any
other species studied, ranging from oval to kidney shaped or
constricted towards one end. The width and length ranged
0.38 x 0,84 to.0.44 ~ 0.94 mm. -Hatching,occurreduaneam
about 10 days at laboratory temperatures.

Larva. - First instar larvae emerged July 7 and were
placed on an immature flowering spike of Phacelta ramosts-
stma which had been taken at Chipmunk Flat six days earlier
and kept in water. Within two hours, several had begun skel-
etonizing leaves. Usually the underside of the leaf was sel-
ected. Later emerging larvae were placed in plastic vials
with leaves and began feeding successfully. By the fourth
day successfully established larvae were still in the first
instar, located either on the underside or in curled portions
on the upperside and had skeletonized several small spots.

After the 9th day larvae were supplied with leaves from
a greenhouse Phacelta ramostsstma transplanted from Chipmunk
Flat on June 24. The cut pieces were accepted but turned
black after a day or two. A leaf of Phacelita dtstans from
San Francisco was supplied to second instar, 11 day larvae,
but little attempt was made to feed on it. Thereafter, the
greenhouse P. ramostsstma was used.

By the 16th day all larvae were third instar. Two days
later some individuals had reached the fourth instar and
were feeding on the fullileaf thickness. Generally, very
little silk was used in visible shelter preparation.

On the 24th day third and fourth instar larvae were
transferred to a branch of P. ramostssitma in water. All be-

Biological studies on Ethmia BHF,

gan feeding without any apparent shelter.

The four surviving larvae reached the fourth and fifth
instars by the 28th day, and Phacelta ramostssitma from Don-
ner Pass was provided.

Some larvae were full grown by the 31st day, appearing
blackish with a dull orange-brown dorsal median band. The
final larva began its cocoon on the 39th day.

Larvae were observed in the field at Donner Pass 30
days after adults had been collected at this site. By this

time the plants had reached full to late bloom in somewhat
sheltered, east facing exposure. The larvae, in the third
and fourth instars, lived in thin webbing shelters among

the inflorescences, usually more or less on the underside
of the flowering spike, rather than between the geminate

flower rows. The silk was not easily visible, but presence
of the larvae was evident by frass, retained by the hairy,
viscid texture of P. ramosissitma. Lower leaves were thor-

oughly investigated and neither larvae nor signs of larval
feeding were found.

Two larvae reached the final instar one day after this
Field collection, and the first larvae spun cocoons on the
12th day following collection (42 days after adults were ob-
served at the site)

There appeared to be five instars, based on unsexed
head capsule measurement (fig. 5). However the size range
in later instars suggests a possible sixth instar in occa-
sional individuals.

Second instar: (None preserved) HC [5] 0.36-0.40 mm,
pale orange with slightly darker mottling.

Third instar: Length 5.0-9.9 mm; HC 0.55-0.78 mm, or-
ange-brown to dark brown, mottled paler; ThSh pale orange-
brown; Pin dark but somewhat diffuse; D whitish, not well
defined; DL scarcely evident, ochreous yellow; AbdCr 7-8;
AnCr 6-9 (usually 8-9).

Fourth instar: Length 10.4-12.0 mm; HC 0.86-0.96 mn,
white anteriorly with dark brown posterior markings; ThSh
mottled laterally only; Pindark, small; D unpigmented, not
well defined; DL pale ochreous or olive-green with whitish
encircling pinacula; AbdCr 9; AnCr 10.

Prpen, the¢ar, Lad: Length 12.2=14 02 mms, AC, 1.22132
mm, white anteriorly with black posterior markings; ThSh
mottling restricted to small areas at posterior margin; D
well defined, unpigmented to dull orange-brown; DL, LV well
defined, dark gray to blackish, mottled with unpigmented
areas; L well defined, pale, unpigmented or tinged with
orange; AbdCr 13-15; AnCr 15-17. Segment AQ with 3 tiny
Secondary setae just anterior of LV seta.

38 nadta

Pupa. - Full grown larvae were placed “in’a Salveutus
with pieces of dry Phacelta stems from Donner Pass. The
stems were 2-4 mm in width and were hollow or had a soft,
pithy context throughout. No larvae used these stems for
pupation. Cocoons were spun in the upper and lower corners
of the tin. Kept at room temperatures and humidity, the
pupae dessicated prior to development. Cocoons were about
12 x 4.5 mm, with a dense, white papery cover and little
internal silk mesh.

Pupae ranged 6.8-7.1 mm in length. The anal legs var-
ted in divergence, with one individual having them nearly
adjacent. The distal portion had no lateral development
and bore about 18 hooked setae. The frail, posterior ''cre-
master'' setae were all short, possibly broken, in the indi-
viduals examined. They originated from a depressed, smooth
area subtending lateral humps.

ETHMIA NADIA CLARKE
Ethmta nadta Clarke, 1950, J. Wash. Acad. Sci., 40:161.

A difficult taxonomic problem exists concerning rela-
tionships of nadia and albistrigella. The present species
occupies generally warmer, drier (Upper Sonoran and Transi-
tion Zone) regions than the boreal sites (Canadian and Hud-
sonian Zone) observed for albistrigella in California. In
addition, it appears that yadtaq is primarily crepuscular.
Possibly ‘it ‘is not obligated to diurnal activity, by Tou
night time temperatures as is albistrtgeliag at higher eleva-
tion stations.

Study areas. - 1) Fowler's Camp, 5 miles east of Me-
Cloud, Siskiyou Co.;1) male at Coleman Lantern July oy poe
2 males, 1 female, net collected, apparently actively flying
in late afternoon and at dusk, July 14, 1962; negative re-
sults in examination of Phaeceltaqa mutabilis, July 21, 1966.
2) Hills back of Citrus Experiment Station, Universimeyuan
California, Riverside, Riverside Co.; larvae on Phaceltia
ramostssima Vat. suffrutescens, May 13, 1962 (62E8). 3)
Herbert Creek, 3 miles west of New Almaden, Santa Clara Co.;
| female net collected between 1:00-2:00 P.M., April 20,
1966 (A.J. Slater and J. Powell)® retained alive (eouemm™

Adult behavior. - Although the moths have been collec-
ted in March and April in southern California and June and
July in northern California, it seems likely that a single
annual flight is involved. lt occurs late in the season
compared to other species with small eyes, and is correla-
ted with flight ‘later in the: day, at’ least Inte a icrepusca-
lar phase. In addition to afternoon, dusk, and evening
collections listed above, single adults have been taken in
the daytime: at Riverside, on flowers of Cryptantha tnterme-
dia (P.H. Timberlake); at Fairview, Tulare County, at midday,
April 27, 1964 (P.A. Rude): and 9 miles’ south of Fairview,

Biological studies on Ethmia Bi

in late afternoon, April 29, 1964 (P.A. Rude).

A reared female was observed in January 1963, over a 16
day period. No males were available. At laboratory temper-
atures this individual was active, crawling and feeding at
damp cotton, at dusk and each evening with lights on in the
room. The moth was not active in morning hours and was not
observed to move much during the afternoon. No eggs were
laid. When prodded during evening activity periods, the
moth would feign death, dropping to the substrate on its
back, with the legs tightly clasped to the body. After a
few minutes activity was resumed.

The female from Santa Clara County (66021), caged in
April, 1966, was observed only on the first night. It was
not active between dusk and 9:30 P.M., or between 5:00 and
5:30 A.M., resting in the quiescent posture. Retained at
outdoor temperatures in a 100 x 85 mm jar with a bouquet of
Phacelia distans, (the only Phacelta discovered at the col-
lection site), the moth lived only five days. At least two
eggs were deposited between 5:30 P.M. and dusk on the first
day and ultimately 24 eggs were laid. Oviposition sites
varied in these conditions (crowding and unnatural orienta-
tion of the Phacelta branch may have been factors). Half
the eggs were placed on leaves, both upper and lower surfa-
ces, with the remainder on flower heads (4), stems (2) and
the nylon screen over the jar (4).

Egg. - The eggs were nearly cylindrical, measuring 0.40
Zeoeeoe to 0.36 x 0.85 mm.

The eggs were placed ina refrigerator from April 25 to
May 3, and were then stored at room temperature. Hatching
occurred after about 17 davs (including the 9 days in cold
storage).

Larva. - Newly hatched larvae were placed on a cut
sprig of Phacelta tanacettfolia from the Botanical Garden in
a salve tin. Two day old larvae had established mostly on
spots under leaves against the salve tin surface. Feeding
occurred as skeletonized spots on either upper or lower sur-
faces of the leaves. However, as the plant material began
drying, by the fifth day, all larvae dispersed and escaped
owing to a faulty container.

Field collected larvae at Riverside in the final and
penultimate instar differed markedly in appearance from
albtstrigella, appearing olive-green with a pale dorsal band.
The plants were in full bloom and larvae spun a thin web
which enclosed a terminal raceme or a leaflet or two.

Penultimate instar: Length 13.0 mm [1]; HC 0.95-1.07
mm [4]; ThSh not differentiated; Pin minute; no integumental
pigment; AbdCr 11-12, uniordinal; AnCr 11.

Final instar: Length 16.5-17.5 mm; HC 1.22-1.28 mn,

Te) semtlugens

orange, strongly mottled with whitish; Pin small, black; D,

L fairly well defined, yellowish; DL pale ollve—-evay eae
tled, with whitish encircling pinacula; AbdCr 17-19, biordin-
al mesally; AnCr 17. Segment A9 with about 12 tiny, unpig—
mented secondary setae in a row between LV and V setae.

Pupa. - Cocoons were spun in the leaf material and in
folds. of paper toweling. The outer layer was dense, white,
tough, paper-like and could be torn when dry. Inside,
cocoons had an ill-defined but strong mesh surrounding the
pupa, making it difficult to extract pupal shells intact.
Pupae were formed before early August. Successful emergence
occurred from nearly all those which had pupated, although
they were stored in laboratory conditions. Emergence took
place in December and January, well ahead of that of field
conditions.

One pupa measured 8.2 mm in length. The anal legs were
irregularly to strongly divergent or curved, with a slight
lateral development distally. About 30 setae were located
in the anchoring group on each leg. The ''cremaster'' setae
were short (possibly broken), in the individuals examined,
and were borne in a shallow V-shaped, roughened depression.

ETHMIA SEMILUGENS (ZELLER)

Psecadta semilugens Zeller, 1872, Verh. Zool.-Bot. Ges.,
Wien, 22:561.

This species is widespread in arid areas from Colorado
to Chihuahua and southern California (Powell, 1959, 1971).
Although there had been only a single record for California,
we were fortunate in discovering larvae on two species of
Phacelia at one locality at the northern end of the Panamint
Valley in 1969.

Study area. - Darwin Wash, 1-3 miles west of Panamint
Springs, Inyo Co.; larvae on Phacelta calthifolta, May eee
1969 (P.A. Opler) (69E65); larvae on P. ealthifoltia and P.
oy a May 14, 1969 (J. Powell and P.A. Rude) (69E78,
7/2) Vee

Adult behavior. - Collection records indicate this spe-
cies is facultatively double-brooded, with flight records
for late February and March to September, but mostly in April
and July. Records in California are for April and May, and
individuals we reared either emerged in July or went into a
prolonged diapause. The moths are nocturnal, judging from
eye size and light attraction records. Adults were not ob-
served in the laboratory.

Egg. - Unknown.

Larva. - Individuals of at least four instars were found
on annual plants along a rocky roadside and wash bottom, a

Biological studies on Ethmia 4]

site which had been heavily eroded during the preceding win-
ter. The caterpillars lived externally on the undersides of
leaves and stems without any visible webbing. Feeding evi-
dently occurred entirely on leaves, although both host spe-
cies were in bloom at the time. In the laboratory, larvae
were housed in polyethylene bags or plastic freezer dishes
and continued to feed more or less exposed. Foliage of the
two plants became mouldy easily. However, larvae did not
Seem to be susceptible to disease outbreak and provided with
refrigerated leaf material, larvae matured 7-14 days follow-
ing collection.

Cocoons were formed in folds of paper toweling. No
soft woody substrate was offered. Mature larvae took on a
pinkish cast while wandering in search of pupation sites.

Head capsule measurements did not clearly define in-
stars, and there may be six instars in this species. The
following diagnosis represents a somewhat arbitrary instar
division, based in part on crotchet numbers and secondary
setae.

Seeond instar (?): [4] Length 3.5-6.0 mm; HC 0.32-0.48
mm, unicoltorous dark brown; integument unpigmented, body
appearing more or less uniform pale green; ThSh brown, near-
ly unicolorous to blotched; Pin dark, conspicuous, but rel-
atively smaller than in later instars; AbdCr 9-13, uniordin-
al, essentially a complete circle; AnCr 7-8.

t
0.75 mm, color of HC, ThSh, and integument as in preceding
instar; AbdCr 8-15 (usually 10-13), partially biordinal;
AnCr 8-12.

Antepenultimate instar: Length 7.0-10.5 mm; HC 0.56-

Penultimate instar: Length 12.0-14.5 mm; HC 0.85-0.98
mm, whitish mottled with extensive blackish areas; integu-
ment color as in preceding instars, except D yellow, DL dar-
ker greenish, L with a yellow blotch on each segment; Pin
darker, larger; AbdCr 13-16; An€r 10-12.

Final instar: Length 15.5-23.0 mm; HC 1.07-1.36 mm,
white with black markings posteriorly; ThSh unpigmented ex-
cept two lateroposterior, variable black patches; D bright
yellow, DL bright green with unpigmented areas around pinac-
ula; L whitish with large yellow blotch on each segment; LV
greenish; Pin large, black; AbdCr 15-28 (usually 20-24), bi-
ordinal; AnCr 20-24, biordinal; segment A9 with 3-6 small
Secondary setae between LV and V setal groups.

Pupa. - Cocoons spun in folds of paper toweling were
flat, oval, with an opaque, white cover; pupation occurred
within 10 days of cocoon construction. All pupae apparently
entered diapause. A few emerged in early July, after 6-7
weeks at laboratory conditions. The remainder did not meta-
morphose; exposure to outdoor shed conditions through the
following year failed to stimulate completion of development

42 arctostaphylella

and emergence. Some appeared to remain viable after 16
months.

Pupae (figs. 8-9) ranged 8.6-9.1 mm in length and were
unusually dorso-ventrally flattened. Each spiracle was foll-
owed posteriorly by a raised area which was subtended ven-
trally by about 50 tiny spicules. The anal legs were moder-
ately to strongly diverging, well separated at the base,
distally without any enlargement. Each had 15-16 hooked se-
tae. Caudally 4 "cremaster'' setae were borne on each of 2
raised areas corresponding to the anal prolegs of the larva,
and these setae were stronger than on other Ethmia examined,
yet still non-functional.

ETHMIA ARCTOSTAPHYLELLA (WALSINGHAM)

Psecadia arctostaphylella Walsingham, 1880, Proc. Zool. Soc.
LONG ous nL OOOO.

Speculation that the name arctostaphylella is a misnomer
and that Ertodictyon is the host of this species (Powell,
1959) has proven to be correct. Ethmia aretostaphylella has
been found closely associated with Hrtodtetyon in various
parts of California on many occasions, while no evidence
that Arctostaphylos is a foodplant has been forthcoming.
Adults fly in late afternoon and at dusk around Ertodictyon
plants, and they can be found resting on the leaves or flush-
ed from foliage during mid day. They have been taken on £,
caltfornicum at many stations in northern California, on £,
trichocolyx var. lanatum in San Diego County, on EE. ecrasst-
foltum in the Santa Rosa Mountain foothills of Riverside
County, and on £. tomentosum in San Luis Obispo and San Ben-
ito Counties. The study areas cited below are only those in
which early stages have been involved, among the many records
for the moth's occurrence on Ertodtetyon.

SEU Viana. a 1) Carson Ridge near Woodacre, Marin Co.;
1 female, 2 larvae, on Ertodiectyon ecaltfornicum May 14, 1960
(60E1). 2) Hills north of Alpine Lake, Marin Co..5 Misiwiee men
E. californtcum, May 28, 1960 (J.M. Burns and J. Powell)
(60E6); adults on &. californicum, April 17, 1961 (C.D. Mac-
Neill and J. Powell), 5 males, 7 females retained alive (61
D3). 3) Three miles west-of Stoneyford, Colusa Coasveume
larvae on #. caltfornteum, May 1, 1961 (61613) ..) 4) em
miles east of Clearlake Oaks, Lake Co.; 1 mature larva on £.
californicum, May 11, 1961 (61£14). 5) Two miles east of
Groveland, Tuolumne Co.; 1 larva on £. caltfornteum, June 12,
1961. 6) Mt. Diablo,. 3000: feet, Contra Costa Co.s, 2ighaeee
on. F., caltforntcum,. July..\75 96) (6162). 7). iE ght ameter
north of Boulder Creek, Santa Cruz Co.:; .l. male,..1 femane cum
cocoon on £. ¢californtcum, Aug. 11, 1962 (62H6). 8) Five
miles east of Boulder Creek, Santa Cruz Co.; adults, cocoams,
larvae on &. ealtfornicum, Aug. 11,.1962 (62H7).

Biological studies on Ethmta 43

Aeault behavior” — lh the foothills of. central Cakiforn-
ia the moths fly as early as February, commonly in April and
in all subsequent months until September. Adults and larvae

occur together during summer, suggesting over!apping genera-
tions. At higher elevation sites spring emergénce occurs in
May, and only two generations may obtain.

In the field the moths become active before sundown and
feyosneo the night, according to light attraction records.
In the laboratory activity began by 6:00 P.M., prior to sun-
See, and Was highest during the next two hours. Some indi-
viduals remained active as late as 11:00 P.M., but they
moved more slowly and activity generally appeared to dimin-
ish late at night, although temperature change was not occur-
PL vg «

When at rest during the daytime, both in captivity and
in the field, the moths assume the characterisitic quiescent
posture and often perch on the upperside of the elongate
Ertoditctyon leaves, oriented with the body axis along the
mid vein. The white and grey color pattern causes the moths
in this position to resemble bird droppings.

Mating pairs were observed four times. One pair was
Swept from Ertodietyon caltfornitcum at Mt. Tamalpais, Marin
County, between 4:00 and 4:30 P.M. on a cool, windy day in
mea Mareh, 1964, by C.W. O'Brien. In the laboratory one
pair was first seen at 8:30 P.M.; the couple moved at least
once, but remained stationary from 9:45 to 11:30 P.M. Sep-
akaeron coecurred ‘between 11:30 P.M. and 7:30 A.M. © The sec-
ond pair apparently mated between 8:00 and 9:30 P.M. and
Femarined in coition until after 10:30 P.M. The other copu-
lation occurred five days after the adults were caged, when
the moths had become very worn appearing. The pair was ob-
served at 7:30 A.M., having mated sometime after 6:30 P.M.

Oviposition occurred at various times of night. One
female was observed probing the nylon screen ceiling with
the ovipositor at 6:00 P.M. Many eggs were deposited between
6:00 and 7:30 P.M. and between 7:30 and 10:30 P.M., and a
few were deposited after 11:30 P.M.

About 200 eggs were produced by 7 females. More than
60% of these were placed on the Eriodictyon; 27% were deposi-
ted on the nylon screen. Of those on the plant 90% were on
leaves, but there was no significant difference in preference
for higher or lower leaves on the stem. Even a lower leaf
which was black with sooty mould (as the lower leaves of &.
ealtfornitcum always are in the field) had 15 eggs. About two
thirds of those on leaves were placed on the upperside, and
all but 8 (of 80) on the uppersides were deposited along the
mid vein (figs. 26-28).

In captivity, males lived 4 to 7 days, females 4 to 8
days, but water was not provided after the fifth day.

hh arctostaphylella

Egg. - (Figs. 26-28) The eggs were elongate, and slight-
ly flattened (slightly wider than thick), measuring about
0.40 x: 0583 tos Ona x, O..,90) mm.

During development the eggs turned pink by the third
day. Hatching occurred after .9.to,..10, days at..laboratonm
temperatures. Eggs of the fall generation were not observed,
and the overwintering stage or stages are unknown.

The eggs proved to be impervious to water. Several de-
posited in a field collection vial were submerged when the
vial was used as a water source in the breeding jar. After
five days the vial was allowed to dry. These eggs all hatch-
ed.on, the: lth to0,.13th day after thei nr adepost tron:

Larva. - First instar larvae were placed in salve tins
with immature terminals of Ertodtetyon caltforntcum. Larvae
tied two leaflets together or spun silk between a leaflet
and the container. Feeding occurred as skeletonizing. By
the sixth day most individuals were still in the first in-
Star. Six and eight day old larvae were placed on #.%eaiz7—
forniteum in water vials. The plant kept well in th ismeemiaus
tion and bloomed, but larvae did not establish well. Appar-
ently they wander considerably even though fresh leaves are
available. Leaves in salve tins did not keep well and lar-
vae had to be transferred every few days.

By the 26th day larvae were in the third instar. Feed-
ing at this stage occurred as skeletonizing on older leaves.
Larvae constructed small silken trackways between leaves.

Some larvae had reached the fourth instar by the 32nd
day. No attempt was made to rear these further owing to
difficulties in keeping the plant, which resulted in fre-
quent exposure of the larvae to mouldy leaves.

Larvae collected in the field were of various stages
from about half grown to mature. Most of these were not pre-
served. Larger larvae typically constructed shelters by
Spinning silk across the upperside of one leaf, pulling its
margins towards the center. In new foliage the leaf margins
were often drawn completely together, forming a tubular shel-
ter, open at both ends. On older leaves which had hardened,
the margins were drawn in only partially, forming a hammock
shaped shelter with a silken mat ceiling, under which the
larva rested, oriented along the midrib. The amount of visi-
ble silk varied, possibly with age of occupancy, and some-
times only a thin layer of silk covered the larva which was
visible from above. Even so, and despite the fact that the
larvae are brightly marked with red and black, their general
light green color rendered them inconspicuous under the silk.
In searching, the silk was usually seen first, and probably
the larvae are thus protected from visual detection by lar-
ger predators.

On one occasion (61E13) several larvae were feeding in

Biological studies on Ethmia AS

inflorescences and immature terminal leaves of £. ealtfornt-
cum. None of these shelters resembled the characteristic
single leaf type observed at other localities. Two or three
young leaves, or sepals and flower parts were tied with silk.
As in other situations, many abandoned shelters were pres-
ent. In the laboratory these larvae seemed to prefer leaves,
which were consumed before the flowers. Ertodtietyon blooms
only tn spring; flower parts and developing seed are not
available to summer feeding larvae.

Final instar larvae of FE. arctostaphylella ranged from
strongly marked, with black longitudinal bands and orange
dorsal spotting, to virtually unpigmented, pale greenish
een feny dark pinacula. Some of the Tatter appeared to be
full grown, and !| had no evidence that any individual devel-
oped from one color phase to the other.

First instar: Length meo—2. 0 mm 5 HC 0.27-0.30 mm,
orange-—brown, ocellar area black; ThSh pale brown; setae and
integument unpigmented.

seeond tnstar: None preserved.

Third instar: Length 6.8 mm [1]; HC 0.62-0.71 mm [2]
Beeuwne- “hos det ined laterally only; D defined, pale; DL -in-
distinct, dark gray; Pin small, not surrounded by pale >
areas; AbdCr 6-7; AnCr 8-9.

Fourth instar: Length 10.4 mm [1]; HC 0.84-0.89 mm
(parasitized) [3], 0.87-1.0 mm [5], dark orange-brown with
regular, posterior darkened areas laterad and mesad on each
epsewousol Vobe; heh defined, mottled dark; Pin large,
gave) Gerined, unpigmented; DL distinet, dark to pale gray,
Pin not defined by pale; L and LV not distinguished, pale
and grayish mottled; Abdcr 9-12; 7 Pner 9-12.

Fifth instar: Length 14.3-19.4 mm [3]; HC 1.29-1.35 mm
(starved) [3], 1.40-1.60 mm, orange-brown lateral and mesal
markings not as well defined as in fourth instar; ThSh un-
Digmented except tiny black pinacula; integumental pigment
ae to well developed, when developed, D welt defined,
pale orange or orange-brown; DL black, well defined with
little pale mottling; L well defined; EAE pale, indistinet;
AbdCr 15-16 to 19-20; Aner*19=?1 , Seement AQ with about 8
secondary setae on LV.

One larva from Mt. Tamalpais (unnumbered collection) ex-
ceeds above limits, representing a possible 6th instar.
Fength 18:4 mm; HE 1.73 mm; AbdCr 21, strongly biordinal;
AnCr 23, biordinal.

Pupa. - Pupation in captivity occurred in folds of paper
toweling and in shelters in foliage similar to those occupied
by larger larvae. Whether these were shelters previously
used for feeding was not ascertained. The general behavior-
al tendency to wander and burrow into soft substrates, known

46 discostrigella

for many other Ethmta, does not seem to be consistently
practiced by EF. arctostaphylella. This is the only New
World species which has been recorded as using foliage for
pupation. Walsingham reared the original specimen from a
cocoon in foliage of Arctostaphylos, suggesting the larvae
wander. In field searches | discovered cocoons of this spe-
cies on the Ertodtetyon at two localities in August, 1962.
Five cocoons with viable pupae were located in tightly fold-
ed leaves, these resembling the typical larval shelters ex-
cept more closely closed over the occupant. The dense,
opaque, white outer layer of silk, covered a thin, loose
silken envelope, which was evident at the ends of the leaf
fold.

One adult was reared from a dry flowering stalk of
Yucca whtpplet, collected near Cajon Pass, San Bernardino
County, in December, 1962. The cocoon was located at the
end of a tunnel several cm in length into the woody cortex,
according to the collector, Eric Jessen.

Development by non-diapausing pupae required 11-13 days,
including cocoon formation (60E6).

Cocoons measured about 14 mm in length and were tough
with dense internal mesh. Pupae ranged 8.5-9.5 mm in length.
The anal legs were broad with slight lateral enlargement,
each distally bearing 27-30 hooked setae which were about
0.14 mm in length. The caudal ''cremaster'' setae were borne
in a shallow, flattened trough; all were short, probably
broken in the individuals observed.

Natural enemies. - Braconid wasps of the genera Apante-
les and Mierogaster were reared from larvae of arcectostaphyl-
ella at four scattered localities. Apanteles (n. sp. #141
of W.R.M. Mason): Alpine Lake (60E6, 3 of the 7 larvae not
preserved). Microgaster (n. sp. #22 of W.R.M. Mason):
Stoneyford (61£3, 1 of 6 larvae); Mt. Diablo (6162, 2 ef 2
larvae); Groveland (unnumbered collection, 1 larva).

The three parasitized larvae at Alpine Lake were still
living when discovered, although each already had a braconid
cocoon alongside it. The mothe larvae crawled slowly if
prodded, but there appeared to be no recent feeding in the
shelters. One of the three was retained alive, and it lived
three days after collection. In each case the bracentaenan.
va had emerged from a hole in the side of the third abdomin-
al segment just below the spiracle of the host.

ETHMIA DISCOSTRIGELLA (CHAMBERS)

Anesychita discostrigella Chambers, 1877, Bull. Geol. Surv.
Terri tories 4 seduce.

This is the most commonly collected species of £thmia in
the New World. The adults are nocturnal and sometimes are

Biological studies on Ethmia 47

attracted to lights in great numbers. Despite its abundance
over a wide range in western North America, until recently
nothing was known of its life history (Powell, 1959). &,
discostrigella and the closely related semitenebrella have
diverged from the typical pattern of the genus and feed on
species of Cercocarpus (Rosaceae). In Great Basin regions

of eastern California, typical dtscostrigella is associated
with Cerecocarpus ledtfoltus. In cismontane parts of the
state, where the moths are generally more bluish white in
appearance and the name subcaerulea Walsingham is applicable,
C. montanus (=betulotdes) is the principle host. For pur-
poses of the present discussion the two are treated together.

The moths have been flushed from foliage of Cercocar-
pus at a number of sites: from C. mtnuttflorus at San Diego,
from C. montanus in the mountains of San Diego County, Kern
County, and Lake County, from C. alntfolta on Santa Cruz
Island, and from C. ledtfoltus in the Warner Mountains, Mo-
doc County. The study areas listed below are those which
have involved the early stages. In addition, M.M. Furniss
@rovchne Uns. ‘Forest Service sent~™me a large series of larvae
and reared adults from Cercocarpus ledtfoltus collected in
Owyee County, Idaho.

Study areas. - 1) Crooked Creek, 10,150 feet, White
Mountains, Mono Co.; young larvae on Cercocarpus ledifoltus,
July 4, 1961 (61G1); 2 females at light, July 22, 1961, re-
tained alive (61G4)._2) Lee Vining Camparound, Mono Co.:
young larvae on C. Ledtfolius, July V9; 1961 (6163). + 3)
Make Pilisbury, Lake Co.; 1 female, at Vight’, April 3, 1962,
retained alive (62D1). 4) miles east of Monitor Pass, Mono
Co.; adults at light, June 30, 1962, 2 males, 4 females re-
tained alive (6263).

Adult behavior. - Collection records from a station in
Monterey County, where a continuous sample of insects attrac-
ted to light was made throughout a season, indicated that
three or more overlapping generations obtain (Powell, 1959).
In Great Basin areas probably a single flight, in June and
July, is normal. Under laboratory conditions pupae result-
ing from eggs laid in June and July in Mono County did not
emerge the same season but underwent diapause, emerging the
following spring.

Although the moths are easily startled into flight dur-
ing the daytime, both in the field and laboratory, even in
early morning, normal activity is nocturnal. Caged females
became active at dusk and engaged in oviposition behavior
then. Whether or not lights were directly on them seemed
not to affect behavior of females except they tended to con-
gregate in the portion of the jar towards the light. The
moths are active all night under favorable temperature con-
ditions, judging. from light attraction records.

Mating was not seen in the laboratory, and field obser-
vations suggest that it may occur only late at night. One

48 dtscostrigella

pair was taken from a congregation of scores of individuals
on a vertical sheet before a 15 watt blacklight, between
12:30-2:00 A.M., east of Monitor Pass, June 25, 1962. Num-
erous mating pairs were observed in tree foliage at Fandango
Pass, Modoc County, in May, 1970, between 9:00-11:00 A.M. by
P.A. Rude.

During oviposition females continuously walked slowly
with the abdomen curled downward and prodded the substrate
with the ovipositor. Females sometimes did this on the
plants provided (Cercocarpus ledifolius for Mono County, C.
montanus for Lake County moths), but more often used the
nylon screen ceiling. I|In one case (62D1) nearly all 28 eggs
were placed on the jar rim under the screen. In the 62G3
lot, four females deposited a total of 82 eggs; only 14 of
these were on the Cercocarpus, 8 of those on the silk of an
abandoned caterpillar shelter. Propensity for selection of
other fibrous and roughened substrates was shown. About 30%
of the eggs were placed on top of the rim of the jar, between
the rim and the appressed nylon ceiling; another 25% were
located on masking tape on the floor of the container (but
only 4 eggs were placed on the smoother cardboard which was
of greater area). Three were placed on a patch of cotton
fibers which had stuck to the vial holding the plant. Of
those on the Cercocarpus two eggs were placed on terminal
stems, adjoining leaf bracts, etc., but none were laid on
the larger, woody stems.

These oviposition sites suggest the possibility of use
of the elongate, twisted, soft-hairy style of the fruit,
which are persistent on the trees, for egg placement in the
field.

Adults did not survive well in captivity, the Lake
County female living 7-8 days, those from Mono County even
fewer.

Egg. - The eggs were somewhat irregular in outline, evi-
dently conforming somewhat to the substrate. Those from one
61G4 female were oval, flattened, tapering in outline toward
one or both ends and measured 0.70 x 1.27 to 0.72 x 1.40 mm.

White when first deposited, the eggs turn bright pink
on the second day, remaining so until just prior to hatching
when the dark larval head capsule becomes visible and the
eggshell looks whitish, semiopaque.

Hatching occurred in 9 days in July, in 10 days in April
at laboratory temperatures.

Larva. - First instar larvae (6263) were placed on Cer-
cocarpus ledtfoltus from Monitor Pass, which had been in
water 10 days. Nine days later some had reached the second
instar. Larvae at this stage were inconspicuous, living in
Crotches of twigs and subsessile leaves, with little visible
Sa ike Feeding occurred as small round skeletonized spots,

Biological studies on Ethmia 49

mostly on undersides and on apical half of leaves. Young
larvae were easily disturbed and quickly dropped down on
silken threads at the slightest stimulus.

Cercocarpus ledtfoltus from the collection site was pro-
vided at 9, 19, and 35 days, after refrigeration, and seemed
to take up water and serve adequately as larval food.

By the 16th day larvae had reached the third instar.
By this time, and thereafter, the larvae were extremely re-
active to external stimuli - prodding caused them to wriggle
backwards extremely quickly, so as to appear to jump, often
going 10 to 20 cm on a flat surface.

At 20 days most individuals were in the fourth instar,
and all larvae had reached the penultimate instar by the
25th day.

By the 35th day all larvae had reached the last instar.
As in the case of £. plagiobothrae, two distinct color
phases were shown: a paler one showing bluish dorsolateral

bands and a lighter orange dorsal band, and a dark form,
which was more common, with the dorsal band yellow-orange to
rust-orange, the dorsolateral bands black.

The final full grown larvae, in the pale state, were
preserved on the 4lst day.

Larvae collected as second instar at Crooked Creek re-
quired a longer period to mature, probably owing to poorer
food conditions. These larvae were provided with cut twigs
Diet. Ledtyjolius in salve tins. After. 13 days fresh C. mon-
tanus from Contra Costa County was provided. All subsequent
feeding took place on this plant. Intermittently the plant
material dried, leaves frozen for 14 days were provided,and
fresh C. montanus was provided again on the 36th day.

The first cocoon was formed 48 days after collection of
second instar larvae, and the last larva died after the 58th
day when additional foodplant from the freezer was added.

There appear to be five instars (fig. 6). The rather
wide spread in head capsule measurements in the final two
instars may have been caused by differential laboratory con-
ditions, since field collected larvae were taken in young
instars.

First instar (62D1): Length 2.3-3.0 mm; HC 0.31-0.30
mm, pale tan, slight brownish spots; Pin visible on thorax.
(61G4): Length 3.1-3.3 mm; HC 0.34-0.38 mm, pale tan with
brown dorsolateral and venterolateral spots; ThSh brownish
laterally; Pin brownish, diminishing on posterior half of
abdomen. (62G3): Length 2.7-3.1 (one day) to 4.2 mm (9
days); HC 0.36-0.39 mm, pale tan becoming darker at 9 days;
Pin pale brownish, becoming darker and well defined on whole
abdomen.

50 dtscostrigella

Later instars are characterized on the basis. of 62G1
and 62G3 specimens.

Second instar: Length 4.7 (Teneral) - 6.0 mm; HC 0.50-
0.55 mm, dark brown without appreciable mottling; Pin dark
brown, those of ThSh larger; integument otherwise without
pigment, setae dark; AbdCr 6; AnCr 8.

Third instar: Length 6.7-8.0 mm; HC 0.60-0.91 mm, yel-
low-tan with faint brownish mottling and dark and frontal
spots; Pin dark, large; D well defined, pale -ordmge ge
heavily mottled, gray to blackish with a paler (less densely
mottled) median streak; L not well defined, LV with almost
no pigment; AbdCr 10-11 (rarely 24); AnCr 11-13.

Fourth instar: Length 9.1-15.1 mm: HC 0.96-1.24 mm,
yellowish with black adfrontal spots;Pin dark; D well de-
fined, orange with blackish spots; DL well defined, Dilackisn,
darker than 3rd instar, obscuring pinacula; L well defined,
pale; LV, almost as dark’ asi DL, Pan only slightly darker;
AbdCr 13-17; AnCr 18-22, biordinal.

Fifth instar: Length 15.9-21.5 mm; HC 1.46-1.68 mn,
orange, mottled darker orange along posterior margins; Pin
dark, relatively smaller than preceding instars; D well de-
fined, dark orange (dark phase) or yellow-orange (pale
phase); DL pale gray or blackish, less densely mottled than
4th instar, densest at D and L margins; L pale, nov Wwertede—
fined; LV mottled. gray, pale to: dark, not well) defsueg,
AbdCr 23-30 (usually 28-30), biordinal mesally; AnCr 26-30,
biordinal. Segments Al, A2, AY with 1-3 small secondary
setae on LV.

Pupa. - Cocoons were constructed in the corners of salve
tins, incorporating a few plant parts. One individual used
a rolled leaf. Pupation occurred within ten days after

starting construction of the cocoon. Those pupating in mid
and late August went into diapause and were housed in the
dark salve tins at laboratory temperatures overwinter. Emer-
gence occurred in late April and early May, probably about a
month ahead of the flight period in Mono County.

Pupae ranged 7.7-8.7 mm in length (61G3). The anal legs
protruded ventrally more strongly than in most other species
and were relatively smaller, widely spaced, and strongly di-
verging, sometimes extending almost directly ventrad and lat-
erad. There were 17-18 (rarely 12) hooked setae on each anal
leg. The caudal ''cremaster'' setae were extremely long and
frail, up to 1.4 mm long (twice as long as the hooked setae
of the anal legs), borne on weakly to well developed lateral
humps. Usually they were broken off in the cocoon.

Biological studies on Ethmta 51

ETHMIA SEMITENEBRELLA DYAR

Peemea semttenebrella Dyar,.1902, Jour. N..Y..Ent...Soc.,
Be 204,

As discussed elsewhere (Powell, 1959) this and the pre-
ceding species, dtscostrtgella are closely related. Subse-
quent studies have shown that the two share similar biolo-
Giles, wsing species ef Cercocarpus as hosts... In the origi-
nal description Dyar mentioned that £. semttenebrella was
reared from ¢, parviflorus in Arizona.

Study apnea. -2Four mi lesveast of Monitor Plass, .-Mono Gox<>s
nest light, June 24 and 30, 1962, 2° males, 6 females
fetaimed,alive,on,latter date (6262).

Adult behavior. - This species is geographically and
ecologically restricted compared to discostrigella, and
semttenebrella appears to have only a single annual flight.

The moths are nocturnal. Adults were caged after 36
hours storage in a field ice box. They became active at
dusk and showed a similar activity pattern to discostrigella
from the same locality. Neither oviposition nor mating was
observed. Females behaved similarly to those of discostri-
fevvautin) prodding the ovipositor through, the nylon screen.
All the semitenebrella died 5-6 days after collection.

Only 44 eggs were deposited by the 6 females. As in
Ever ecese Of diseostrigella, a preference for roughened sur-
faces was shown, but 30 of the 44 were deposited on the
f1een Of the jar, possibly in,.part.a function of age or
weakening of the females. All but 3 of these were laid on
fiasking) tape or in:a crease in: the cardboard. Only 3: eggs
Weme deposited on the Cereocarpus.. Seven eggs were placed
On the nylon screen, but none were at the rim of the jar
adjacent to or under the appressed nylon, a site used for
30% of discostrtgella eggs.

Egg.a.- AS, in dzrseostrtgetia, eggs, of ;the present Species
were not regularly rectangulate or ovate, but varied to
some extent with the substrate, often tapering towards one
end. Those deposited on the nylon screen measured 0.66 x
heo,to. 0.70 x 1.32 mm.

MA scupned uniform dull reddish prior to the fttth day,
darkening only shortly before eclosion. Eggs began hatching
Hive ia, about .10.5-11 days after oviposition. | Most were
transported on a field trip July 13-15; and those remaining
unhatched survived transit in an uncooled car at 40° C air
temperature, conditions which were lethal to young larvae,
hatching between 8:00 and 11:00 A.M. the following day, about
badays vincubat.ton.

Larva. - First instar larvae were placed on terminals of
Cereocarpus leditfoltus which had been in refrigeration 12

52 semttenebrella

days. Those emerging prior to July 15 did not survive auto-

mobile transport in 40° C air temperature. The remaining
first instars hatched July 16 and were placed on 15 day old
refrigerated C. ledifoltus. They showed a marked tendency

for positive phototropism during daylight hours, crawling to
the top of a vial, away from leaves or to the side of a
branch toward the light.

Larvae established in small webs in crotches of leaves
and twigs or between leaves. Larvae had reached the second
instar before the l4th day. Additional C. Ledifolius from
Monitor Pass was provided on the 14th day, after 30 days in
refrigeration.

The third instar was reached on the 15th to 16th day,
and the fourth by the 20th day. -Although the orlg@inateeraneh
had become covered with fine mold by 10-14 days, larvae had
not left it and were transferred to the fresher foliage
which had been added to the bouquet 6 days previously. At
this time evidence of larvae had become quite noticeable,
with considerable webbing which at times collected frass on
the uppersides of leaves.

The final instar was reached by the 29th day and the

last full grown larva was preserved on the 35th day. To the
unaided eye mature larvae appeared steei blue-gray with
bright yellow dorsal and lateral bands. The venter was

bright pink, differing from dtscostrigella which had a pale
venter.

Too few specimens were preserved to enable precise de-
termination of the number of instars.

First instar: Length 3.3-3.5 mm; HC 0.45-0.47 mm, pale
tan with black ocellar area, becoming entirely dark Drown au
maturity. No integumental pigment.

Seeond tnstar: Length 8.2 mm [1]; HC 0. 64-060 mma,
pale brown, mottled darker; Pin brown, rather large? anresn—
ment otherwise unpigmented, setae black; AbdCr 8-11 seanmite TL

Penultimate instar (none preserved): Length 9-10 mm; HC
1.04-1.28 mm [2], pale orange, shaded brownish, not strongly
moveled, DP yew lonarsla sy Ula eenic ws lamin,

Final instar [3]: Length 15.9-23.0 mm; HC 1.70-1.87 mm,
Orange, mottled with white laterally; Pin dark ®orowm, seaqee—
ly differentiated from integumental dark areas, strongly con-
trasting in pale areas; D and E distinct yellow, Uirear:
steel gray to blue-gray, irregularly mottled with small white
Spots; LV only lightly mottled; V bright pink, a subintegu—
mental color; AbdCr 28-30 or 32-34: AnCr 30-34.° Segment Ag
with 12-14 small secondary setae on a sclerotized patch ex-
tending 1/2 the distance to V seta.

Biological studies on Ethmia 53

ETHMIA TIMBERLAKEI POWELL

Ethmia timberlaket Powell, 1971, U. Calif. Publ. Ent.; in
press.

This species is closely related to geranella Barnes and
Busck, and may prove to represent a segregate of that species
when more is known of both. These and related species dif-
fer from most other Ethmia in life cycle, feeding as larvae
in spring, aestivating as pupae, and flying in fall. The
biology of FE. macelhostella Busck, a member of the group in
eastern United States, was studied by Busck and Heinrich

Gh922).

Study area. - Hill back of Citrus Experiment Station,
University of California, Riverside; larvae on Phacelia ram-
ostsstma var. suffrutescens, March 21 and 24, 1961 (P.H.
Timberlake, R.L. Langston and J. Powell) (61C12, C13); lar-
vae on P. ramosisstima, May 13, 1962 (62E7).

Adult Behavior. - Only a single individual has been
field collected, at light between 8:30 and 10:00 P.M. on
October 17, 1960, near Desert Springs, San Bernardino County.
Closely related species have been taken at lights between
mid September (8000 feet elevation) and November (1000 feet).
Adults of E. timberlaket emerged prior to November 8 (61C12)
and between November 1 and 19 (62E7).

Reared moths were observed in late November, 1962. One
male and two females already in worn condition were caged in
a dry jar with debris and cocoons from the original rearing
container. The moths were inactive during daylight and
appeared reluctant to move at night with lights on in the
room. At this time they moved only by short, quick ''jumps'"'
when disturbed by the observer. They were active at night
with the lights off.

Fertile eggs were deposited prior to November 19, but
not after that date. Although no water was available, the
moths lived 11-14 days after first observed, when already in
worn condition.

Numerous eggs were deposited, primarily in aggregated
groups, not in any systematic arrangement, around the glass
side of the jar near the upper rim. A few scattered eggs
were laid on the dry Phacelta foliage and paper toweling.
Most had turned reddish by November 19 and apparently en-
tered diapause.

Although kept in a dry container at laboratory tempera-
tures, about half of the eggs hatched at sporadic intervals
during the following spring.

Larva. - In late March, 1961, larvae of at least the
final three instars were present on Phacelta ramostssima.
Most were in the last instar. In mid May, 1962, only full

54 timberlaket

grown larvae were present, and evidences of feeding indica-
ted that most had already left the plants.

Larvae of £. ttmberlaket differed in habits from most
other species studied (including nadia at the same site) by
living exposed on the leaves, without any shelter. During
the daytime most were concentrated towards the lower portions
of the dense foodplant clumps, rather than exposed in direct
Sunlight. In several cases the bushes grew adjacent to
large boulders on a southerly exposure. The caterpillars
rested toward the back of the clump, in the shade, where the
foliage was most dense. Almost all perched on the under-
sides of stems or main midrib of the compound leaves.

Presence of larvae was evidenced by scattered frass
toward the distal end of the branch, evidently held by the
viscid hairiness of the plant. Probably larvae moved out-
ward at night to feed, and rested under the stems inactively
during the day.

A few small webs were found on undersides of leaves,
with associated head capsules. lt is assumed these were
moulting webs, but it may be that early instars, which were
not observed, construct weak shelters.

In rearing, larvae were housed in 85 x 100 mm jars in
field conditions for 7 to 10: days and severe moisture con-
densation and moulding of foodplant resulted. However, no
disease symptoms developed. Fresh Phacelta, presumed to be
ramostssima, from San Diego County, was provided on the 5th
day,.but littie,or sno, feeding ocecurned on its

Most of the larvae successfully formed cocoons by the
12th day after collection.

Second instar (?) [2]: Length 6.0 mm; HC 0.47 mm, en-
tirely ‘dark brown; Pin tiny, dark; almost no | Gtheg yume oan
mental color; D, L weakly white, Abder 8-10; AnCr 7.

Third inatar (?);. length 6.0-7.8 mm (venereal toes a0
mm; HC 0.61-0.68 mm (teneral and parasitized), 0.73-0.83 mm,
dark brown with pale mottling anteriorly; D defined, Watersng
DL pale gray; Pin black,small; integument otherwise unpig-
mented; AbdCr B=ills Antic 7-1) (usualy MO—m

Penultimate instar: Length 10.4-12.0 mm; HC 0.87-0.97
mm, dark brown posteriorly, whitish anteriorly; D well de-
fined, yellowish; DL mottled grayish (appearing bluish green
in life); L distinct, yellowish; Pin dark, small; Apdemee =
AGO MNinloae Walrad 6 ren

Final instar: Length 9.5 mm (unfed?), 15-6-117 .2)mmagne
1.0-1.23 mm, white, mottled with brownish black posteriorly;
ThSh defined by small blackish mottling; Rin small , hack a!
yellow; almost no other integumental color, DL pale grayish,
lightly mottled (appearing pale bluish green in life), leav-

Biological studies on Ethmia 55

ing irregular unpigmented areas around pinacula; AbdCr. 14-20
Ceeuatiy 17-20); AnCr 16-21.

Pupa. - Cocoon formation occurred mainly in folds of
paper toweling in the rearing container; a few were formed in
mouldy foliage. Pupae were formed soon after cocoon construc-
tion. At least one pupa was present by April 5, 12 days
after the larvae were collected.

In the field no cocoons were located on the foliage,
even in late season condition, in May, 1962. The closely re-
lated HE. macethostella was reported to burrow into bark of
trees and logs for pupation (Busck and Heinrich, 1922). At
Riverside, the dry chaparral association contains no plants
with large woody trunks and appreciable thickness of bark.
The pithy, dry stems of previous years! Phacelta growth was
the only likely site evident in which larvae might burrow,
but search of a large random sample of preceding years'
stems revealed no abandoned cocoons.

The cocoons were about 11-12 mm long and exteriorly were
papyrus-like in consistency, not translucent and could be
torn like paper. Inside there was little loose mesh, and pu-
pal shells could be easily extracted without breakage.

Pupae measured 7.5-8.2 mm in length. The cremaster se-
tae were observed intact on several individuals, about 0.11]
mm in length, very frail, curving towards the tip. The anal
legs were widely spaced (0.33 mm apart at base), not diverg-
ing, and short, the free part only about 0.23 mm in length.
The distal end bore 18-20 anchor setae in several examples.

Cocoons were stored in dry jars at room temperature and
successful emergence occurred from nearly all, during a three
week period in late October and the beginning of November.

Natural enemies. - The colony at Riverside was affected
by a braconid, an undescribed species of Microgaster (n. sp.
#8 of W.R.M. Mason). In the laboratory Ethmta larvae reach-

ed the final instar prior to emergence of the braconid lar-
vae. Numerous wasps were reared, and it is assumed that

each affected the host solitarily. However, no estimate of
the proportion of the sample which was parasitized was made.
Under laboratory conditions Mitcrogaster adults emerged in May,
apparently out of phase with any available stage of the
ethmiid. ;

56

LITERATURE CITED

Braun, A.F., 1921. Two weeks collecting in Glacier National
Park. .Brec. Acad. Nat. Seb.) Pht ast: 3352 bee

Busck, A. and C. Heinrich, 1922. Life history of Ethmia
macelhostella Busck. Proc. Ent. Soc. Wash., 24(1):1-9.

Cuming, F.G., 1941. The distribution, life history, and ec-
Oonomic importance of the winter moth, Operophtera bru-
mata (L.) (Lepidoptera, Geometridae) in Nova Scotia.
Canad. Ents ¥/92(6)2135-148-

Dyar, H.G., 1902. A review of the genus Ethmta with des-
criptions of new species. J. New York Ent. Soc.,
10:202-208.

Keifer, H.H., 1936. California Microlepidoptera Vill. Bull.
So. €attt. Acad: Sci. ;,. 3529-29;

Lawrence, J.F. and J.A. Powell, 1969. Host relationships in
North American fungus feeding moths (Lepidoptera:0eco-
phoridae, Oinophilidae, Tineidae). Bull. Mus. Comp.
Zool., Harvard, 138:29-51.

Parker, F.D. and R.M. Bohart, 1966. Host-parasite associa-
tions in some twig nesting Hymenoptera from North Amer-
ica. Pan-Pacific Ent., 42(2):91-98.

Powell, J.A., 1959. Studies on the genus Ethmta Huebner in
western North America (Lepidoptera:Gelechioidea). Was-
mann J. Biol., 17(1):133-151.

1962. Some observations on the minimum temper-
ature threshold of moth activity at light. Presented
at 12th Ann. Meeting Pacific Slope Br., Lepidopterists'
Soc., Santa Barbara [unpublished].

1964. Biological and taxonomic studies on tor-
tricine moths, with reference to the species in Califor-
nia (Lepidoptera:Tortricidae). U. Calif.. Publ Entue
32, 318pp.

1971. Systematic monograph of New World eth-
miid moths (Lepidoptera:Gelechioidea). U. Calif. Publ.
Ent, in’ press’.

Host PLANT INDEX

Boraginaceae

Amsinckia intermedia F.&.hM.

A. lunarits MacBride

A. spectabilis F.€.M.

PemmesselLlaba Gray... -. « « « « «= =
Cryptantha ctreumcissa (H.&A.) Johnst.
C. intermedia (Gray) Beene o2m8
Plagtobothrys nothofulvus (Gray) Bnae
moeteneiius (NWEE-) “Gray. <° S's aes 3

Hydrophyllaceae

Ertodtetyon caltfornicum (H.&A.) Greene

Nemophita maculata Benth. :
Meumemaresrs Wo & A.W 295° Be Gee x
Phacelia cattfornica Cham.
calthtfolza Brand

erenulata Torrey

distans var. australis pone
ramostssima Dougl.

bal seh serfs ly bss]

tanacetifotiia Benth.

Rosaceae

Cereocarpus ledifoltus Nutt.
C. montanus Raf.
C. parviflorus Wooten

Scrophulariaceae

Collinsia heterophylla Buist.

distans Benth. .. :bis6

ramostsstima var. suf -Prutescens Parry

57

-t3

58

EXPLANATION OF FIGURES

Figs. 1-6, larval head capsule measurements in six spe-
cies of Ethmia, Each square represents one individual.
Diagonal-lined squares represent larvae reared from eggs in
lab; solid, half-solid, and shaded squares represent field
collected larvae. Rearing lot numbers refer to data given
in text. Size scale (mm) fis the same in figures aeaem

|. E. plagtobothrae Powell

2. E. albttogata Walsingham

3.. H£.(b. previstriga, Clarke

4s EL segtila rowel)

5. E. a. albistrigella (Walsingham); dotted line indi-
cates hypothetical size of first instar.

6. E. dtseostrigella (Chambers)

Vi 62c2 ~=—*W 69Ds8

Bcavio {| ows
Z Ree — al

59

]
4 68B178
§ 63E1
lt VA) A wall >) “i
2
y 61D2
| ae a
ZA o | 4 |
3
4 69C90

ee ok UU

ae ee. ee ee oe z
0.5

mm 1.0
y 62G3
N 61G3
B ict
V a Sg eee BE cyetae
mm 0.5 1.0 1.5

4

YZ 62en
B cons

iS

6

60

EXPLANATION OF FIGURES

Fig. 7, final instar larva of Ethmta charybdts Powell:
head and thoracic segments I-II, dorsolateral aspect; ab-
dominal segments 6-10, lateral aspect. Body regions: D =
dorsal, DL = dorsolateral,L = lateral, LV = lateroventral.

Figs. 8, 9, pupa of EF. semitlugens (Zeller); 8, ventral
aspect; 9, lateral aspect.

Fig..10, pupa of #. seylta Powell, ventral aspecm.

Figs. 11, 12, e€gg of EF. “eoqutltettetla Buseck, Isiimeee
ting placement on nylon mesh, a substrate commonly selected
by females of various species under cage conditions; ll,
ventral aspect; 12, lateral aspect. Length of egg = approxi-
mately 0.80 mm.

61

62

EXPLANATION OF FIGURES

Figs. 13-20, eggs of Ethmta (approximate magnification
indicated in parentheses)

13, 14, eggs of FE. seylla Powell (69C90) in petiole
axils of Collinsta heterophylla (4x)

15-17, scanning electron micrographs of #&. seytla eggs
and detail of. chorion structure (69C€90); 15, (60x)e tee
(300x) #5 173° (1200x )2

18-20, eggs of EZ. minuta Powell (63D18) in unopened,
hispid inflorescences of Cryptantha intermedia; 18, (12.5x);
VS SCT RICH 20 eae

63

Sees

Ss
.

64

EXPLANATION OF FIGURES

Figs. 21-24, eggs of Ethmia on natural plant substrates
(approximate magnification indicated in parentheses

21-23, eggs of Ey. Db. brevistriga Clarke (6102) a7 ion
sand-encrusted lower branch of Phacelta distans (7x); 22, on
underside of mid-vein of P. dtstans compound leaf (7x); 23,

Siame. .egigs. ( lh2ssaoxKy) +

(62C2) on underside
host which was
larvae

24, eggs of H. plagtobothrae Powell
of Phacelta caltforntca leaf, an abnormal
partially accepted by females but not accepted by

(P55) 8

65

66

EXPLANATION OF FIGURES

Figs. 25-29, eggs of Ethmta on natural plant substrates
(approximate magnification indicated in parentheses).

25, eggs of FE. plagtobothrae Powell (62C2) on setose
stem of Phacelta. caltforntca, an abnormal host (13.5x).

26-28, eggs of FE. arctostaphylella (Walsingham) (61D3)
on Ertodtctyon caltforntcum; 26, on lower branch encrusted
with sooty-mold (7.5x) (the lower foliage commonly becomes
covered with sooty-mold owing to glandular secretions of
this plant); 27; on mid-vein, upperside of leaf (7.5x); 28,
same eggs (12x).

67

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